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BULLETIN OF 

THE BRITISH MUSEUM 

(NATURAL HISTORY) 



ZOOLOGY 
VOL IV 

1956— 1957 



PRINTED BY ORDER OF THE TRUSTEES OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

LONDON: 1957 



DATES OF PUBLICATION OF THE PARTS 


No. i. 


27 March 1956 


No. 2. 


17 August 1956 


No. 3. 


17 August 1956 


No. 4 


28 September 1956 


No. 5 


23 November 1956 


No. 6. 


23 November 1956 


No. 7. 


26 February 1957 


No. 8. 


6 December 1956 


No. 9. 


18 January 1957 



PRINTED IN 

GREAT BRITAIN 

AT THE 

BARTHOLOMEW PRESS 

DORKING 

BY 

ADLARD AND SON, LTD. 



CONTENTS 



ZOOLOGY VOLUME 4 

No. I. British Mites of the subfamily Macrochelinae Tragardh (Gamasina- 

Macrochelidae) . By G. owen evans and E. browning (Pis. 1-4) 1 

No. 2. The evolution of Ratites. By sir gavin de beer (Pis. 5-9) 57 

No. 3. Studies on the Trichiuroid Fishes — 3. A preliminary revision of the 

family Trichiuridae. By denys w. tucker (PL 10) 73 

No. 4. Variation, relationships and evolution in the Pachycephala pectoralis 

superspecies. By ian c. j. galbraith 131 

No. 5. A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae) 
Part 1 : H. obliquidens Hilgend., H. nigricans (Blgr.), H. nuchi- 
squamulatus (Hilgend.) and H. lividus, sp. n. By p. h. greenwood 223 

No. 6. A systematic revision of the fishes of the Teleost family Carapidae 
(Percomorphi, Blennioidea) , with descriptions of two new species. 

By D. C. ARNOLD 245 

No. 7. Variation in the western Zosteropidae (Aves). By r. e. moreau 309 

No. 8. Sagitta planctonis and related forms, by p. m. david (PL 11) 435 

No. 9. Evolutionary trends in the classification of Capitate hydroids and 

medusae. By william j. rees (Pis. 12-13) 453 

Index to Volume 4 535 



INDEX TO VOL. IV 



The page numbers of the principal references and the new taxonomic names are printed in 

Clarendon type. 



466, 48 



abyssinica, Zosterops 

Acarus . 

Acaulidae 

Acaulis 

Acauloidea ..... 

aciculatus, Nothrholaspis 

aciculatus, Macrocheles (Nothrholaspis) 

acus, Carapus .... 

Aepyornis ..... 

aldabrensis, Zosterops maderaspatana 

alfurorum, Pachycephala macrorhyncha 

allmani, Hydractinia 

ambigua, Pachycephala pectoralis 

Amphinema . 

Andrholaspis 

anglicus, Coprholaspis 

Annulella 

Anthomedusae 

anzac, Assurger 

Aphanopodinae 

Aphanopus . 

Apteryx 

Archaeopteryx 

arctica, Cyanea 

arenosa, Aselomaris 

Areolaspinae 

Arum . 

Aselomaris 

Assurger 

Asyncoryne . 

Asyncorynidae 

Athecata 

atromaculata, Pachycephala pectoralis 

aurantiiventris, Pachycephala pectoralis . 198 

aurata, Euphysa 

460, 466, 486, 492, 505, 507, 519, 521 



• 314 
1, 12, 38 

453-534 

1, 509 

523 

46, 48 

. 48 

247-265 

• 64 

• 403 
. 208 

476, 478 

206 

. 481 

4 

4, 16,53.54 

• 5°6 
. 520 

107 

77 
.81-85 

• 63 
60-68 

• 483 

• 476 
9 

509 

481 

106 

459, 509 

• 524 

• 520 
206 



467 
476 



badius, Holostaspis 

balim, Pachycephala pectoralis 

banksiana, Pachycephala pectoralis 

bartoni, Pachycephala soror 

bella, Pachycephala pectoralis 

Benthodesmus 

bermudensis, Carapus 

birpex, Carapus 

blondina, Lizzia 

boraborensis, Carapus 

borbonica, Zosterops borbonica 

Boreohydra .... 



16 
200 
208 
196 
198 
85-90 
270, 272 
265 
481 
279 
403 
507 



Boreohydrinae . . . . -521 

Bougainvillia . .... 476, 481 

bougainvillei, Pachycephala pectoralis . 197 

Branchiocerianthinae . . . .522 

Branchiocerianthus . . . -472 

britannica, Bougainvillia . . .481 

brunneipectus, Pachycephala pectoralis . 208 

buruensis, Pachycephala pectoralis . .202 



12, 



caledonica, Pachycephala pectoralis 

calliope, Pachycephala pectoralis 

caninus, Carapus . 

capensis, Myriothela 

capillare, Eudendrium 

capillata, Cyanea . 

Capitata 

Carapus 

carbo, Aphanopus 

carinatus, Gamasus 

carinata, Nothrholaspis 

carinatus, Macrocheles 

carnea, Podocoryne 

castaneus, Macrocheles 

Casuarius 

caudatus, Lepidopus 

centralis, Pachycephala pectoralis . 

chlorura, Pachycephala pectoralis 

christophori, Pachycephala pectoralis 

cinereus, Carapus .... 

cinnamomea, Pachycephala pectoralis 

citreogaster, Pachycephala pectoralis 

Cladocoryne 

Cladocorynidae 



462, 464, 51 



467, 487, 505 



Cladonema 

Cladonemidae 

Clava . 

clio, Pachycephala pectoralis 

cocksi, Arum 

cognatus, Macrocheles 

collaris, Pachycephala pectoralis 

comorensis, Zosterops maderaspatana 

conferta, Dicoryne 

contempta, Pachycephala pectoralis 

conybearei, Heterocordyle 

Coprholaspis . . . 4, 7, 10 

Cordylophora 

cornucopiae, Merona 

Corymorpha 

Corymorphidae 



202 
. 282 

. 468 
. 476 

• 483 

520 

247-302 

83 

25 

25 

25, 26, 34, 36, 53 

• 476, 478 

42 

63 

93 

198 

201 

198 

295 

197 

203 

511 

524 

519 

525 

476 

202 

509 

28 
203 

403 
476 
201 
476 

53, 54 

476 

476 

462-508 

453-534 



16 



INDEX 



537 



Corymorphinae 

Corynidae 

Corynoidea 

costata, Zanclea 

cristatus, Tentoriceps 

crocea, Tubularia . 

cucullata, Pachycephala pectoralis 

curvirostris, Zosterops curvirostris 

cuspis, Carapus 

Cyanea .... 

cyclopum, Pachycephala schlegelii 

Cyrtocheles .... 

Cytaeis .... 



• 521 
453-534 

. 524 

474. 499, 512, 519, 526 

no 

. 481 

201 

• 403 
. 267 

• 483 

• i95 
4 

• 476 



dahli, Pachycephala pectoralis . 

dammeriana, Pachycephala pectoralis 

decoloratus, Gamasus 

decoloratus, Macrocheles 

Dendrocoryne 

dentatus, Echiodon 

dentatus, Macrholaspis 

dichotoma, Eleutheria 

Dicoryne 

dinema, Amphinema 

Dinornis 

Diplospinus . 



Dipurena 
Dissoloncha . 
domestica, Musca 
Dromaeus 
drummondii, Echiodon 
dubius, Carapus 
dumortieri, Ectopleura 



199 

202 

32 

12, 32, 33, 34, 53 

• 475. 5i3 
. 292 
. 4, 46, 48, 51, 53 
500, 512, 519, 525 
. 476 
. 481 
. 64 
78-81 
460, 475, 493, 498 
4, 10 
13 
63 
288 
270 
491, 496, 509, 519 



475= 



echinata, Hydr actinia 
Echiodon .... 
Ectopleura .... 
efatensis, Pachycephala pectoralis 
Eleutheria .... 
Eleutheriidae 
elongatus, Benthodesmus 
Encheliophis . 

Eudendrium . 
Euphysa . 
Euphysinae . 
eurycricota, Zosterops 
everetti, Pachycephala pectoralis 
eximia, Sarsia . 

Eupleurogrammus . 
Evoxymetopon . 

far eta, Euphysa . 

feminina, Pachycephala pectoralis . 

fergussonis, Pachycephala pectoralis 

filiformis, Staurocoryne . 

fimicola, Nothrholaspis 

flavifrons, Pachycephala 

floccosa, Cladocoryne 



47°, 479, 481 

288-294 

491, 509 

. 208 



500, 



512, 519 

• 525 

88 
295-298 

• 476 
460-507 

• 52i 

• 363 
196 

• 519 
102 

• 97 

. 486 

. 198 

207 

462 

36,38 

136-222 

5ii, 524 



Formica ..... 

fulginosa, Pachycephala pectoralis . 
fulviventris, Pachycephala pectoralis 
fulvotincta, Pachycephala pectoralis 
fuscoflava, Pachycephala pectoralis . 
fusiformis, Rhizogeton 

Gamasina-Veigaiaidae 

Gamasus . . 13, 16, 25, 30, 32, 

Garveia 

Geholaspis . . . 4-10, 

gemmifera, Sarsia 

Gempylina 

gigantea, Monocoryne . 488, 

gilolonis, Pachycephala pectoralis 

glaber, Macrocheles 4, 10, n, 16, 

glabra, Holostaspis 

glacialis, Corymorpha 

gladiator, Macrocheles 

glaucura, Pachycephala pectoralis 

gloriosus Macrocheles (Nothrholaspis) 

goodsoni, Pachycephala pectoralis . 

Gotoea ..... 

gracilis, Diplospinus 

gracilis, Encheliophis (jordanicus) . 

gracilis, Solanderia 

graeffi, Pachycephala pectoralis 

groenlandica, Corymorpha 



13 
200 
196 
196 
202 
476, 478 



42, 46, 48 
. 476 

41-43, 53 
493, 519 
121-127 

509, 523 
206 

17, 19, 53 
16 
496 

38,53 

200 

28 

203 

493 
81 
299 
526 
198 
496 



36 



haeckeli, Margelopsis . . 482, 508, 522 

haesitata, Zosterops curvirostris . . 403 

Halocordyle .... 460, 474, 510 

Halocordylidae . . . . .524 

halterata, Dipurena . . . 475, 493 

hancocki, Encheliophis . . . 298 

Haplochromis .... 225-243 

Hesperornis . . . . . -67 

Heterocordyle ..... 476 

Holostaspella . . 5, 7, 9, 10, 51-53 

Holostaspis . . . 13, 27, 38, 44, 48, 51 

Holostaspis (Holostaspella) . . 51 

Holostaspis Pisentii . . . 13 

Holaspulus ...... 9 

homei, Carapus ..... 273 

hooperi, Stylactis . . . . .481 

houlti, Carapus ..... 279 

hulli, Macrocheles ..... 53 

hulli, Nothrholaspis . . . -25 

Hybocodon . . . . . 491 

Hydractinia .... 475, 476, 481 

Hydrocoryne .... 500, 512 

Hydrocorynidae . . . . -525 

hypochthonius, Macrocheles ... 25 
Hypolytus ..... 460, 506 

Ichthyornis . . . . .67 

imperator, Branchiocerianthus . . 472 

indivisa, Tubularia. . . . 496, 522 

inkermanika, Ostroumovia . . . 483 



538 



insignitus, Macrocheles . . . n, 21, 53 

intermedius, Eupleurogrammus . .103 

japonica, Cytaeis ..... 476 

javana, Pachycephala pectoralis . .196 

jordani, Encheliophis .... 298 

Jordanicus ...... 299 

jubilarii, Pachycephala pectoralis . . 206 

kagoshimanus, Carapus .... 278 

kandavensis, Pachycephala pectoralis . 204 
kirki, Zosterops senegalensis . . . 403 

klossi, Pachycephala soror . . .196 

lacustris, Cordylophora . . . .476 

lamarcki, Cyanea .... 483 

larynx, Tubularia ..... 496 

lateralis, Zosterops . . . -316 

lauana, Pachycephala pectoralis . . 204 

Lepidopodinae ..... 89 

Lepidopus ..... .90-97 

Lepturacanthus . . . . .119 

lepturus, Trichiurus . . . .114 

Leuckartiara . . . . .476 

linearis, Bougainvillia .... 476 

littayei, Pachycephala pectoralis . .201 

lividus, Haplochromis .... 232 

Lizzia ....... 481 

Longicheles . . . . 4, 42, 44 

longispinosus, Gamasus . 
longispinosus, Geholaspis, (Geholaspis) 
longispinosus, Holostaspis 
longulus, Geholaspis (Longicheles) . 
longulus, Holostaspis 



42 
42 
42 
44 

44 



Macrholaspis . . 4, 5, 7, 10, 46-50 

Macrocheles ...... 4-53 

Macrocheles (Coprholaspis) . . .16 

Macrocheles (Monoplites) . . .16,41 

Macrocheles (Nothrholaspis) . . 41, 48 

Macrochelidae . . . . 4, 9, 53 

Macrochelinae . . . 4» 9. 5 1 . 53 

macrorhyncha, Pachycephala pectoralis . 202 
maderaspatana, Zosterops maderaspatana 403 
Malacirops ..... 309-433 

mandibularis, Geholaspis (Longicheles) 

44, 45, 47. 53 



mandibularis, Holostaspis 




44 


margaritiferae, Carapus (Onuxodon) 


. 285 


Margelopsidae 


453-534 


Margelopsinae 




• 522 


Margelopsis 




482, 508 


marginatus, Acarus 




11, 38 


marginatus, Holostaspis . 




19, 36, 38 


marginatus, Macrocheles 




4, 16 


marri, Sagitta 




. 443 


matrius, Macrocheles 


12 


32, 33, 36 


matrius, Nothrholaspis . 




34 


mauritiana, Zosterops borbonica 


• 4°3 


mayottensis, Zosterops semiflava 


• 403 


mediterranea, Porpita 




• 5i9 



melanonota, Pachycephala pectoralis 

melanops, Pachycephala pectoralis . 

melanoptera, Pachycephala pectoralis 

melanura, Pachycephala pectoralis . 

mentalis, Pachycephala pectoralis 

merdarius, Holostaspis 

merdarius, Macrocheles 

Merona 

merula, Saxicola 

michaeli, Aselomaris 

minimus, Macrocheles 

mirabilis, Pelagohydra . . 470 

misakinensis, Dendrocoryne 

misimae, Pachycephala pectoralis 

miurensis, Hydrocoryne . . 500, 51 

modesta, Zosterops 

Monocoryne . 

Monocoryninae 

Monoplites 

montana, Nothrholaspis 

montanus, Macrocheles . . 23, 24 

mouroniensis, Zosterops . 

multistriatus, Diplospinus 

Musca .... 

muscaedomesticae, Macrocheles 

muscoides, Bougainvillia. 

muticus, Eupleurogrammus 

myersi, Cladonema. 

Myriothela .... 467 

Myriothelidae 

Myriothelinae 



197 
204 
198 
199 
197 
21 
21, 22 
476 
209 
481 

44. 53 
, 508, 522 

513 
209 

, 525 

403 

488, 509 

523 
10, 16 

23 

25. 53 

403 

79 

13 

12, 13 

476 

105 

464 

468, 487 

453-534 

• 523 

nemoralis, Macrocheles (Nothrholaspis) . 41 
Neoholaspis ...... 4 

nigricans, Haplochromis . . . 237 

Nothrholaspis . . . . 10, 46, 53 

nuchisquamulatus, Haplochromis . .241 

nutans, Corymorpha . 460-5O8, 519, 521 

nutans, Garveia . . . . .476 

nutricula, Turritopsis . . . 476, 478 



Obelia ...... 

obiensis, Pachycephala pectoralis 

obliquidens, Haplochromis 

obscurior, Pachycephala schlegelii . 

obvoluta, Hypolytus 

occidentalis Macrocheles (Nothrholaspis) 

octona, Leuckartiara 

octopunctata, Rathkea 

Onuxodon ..... 

opacus, Gamasus .... 

opacus, Macrholaspis 

ophiogaster, Dipurena 

orioloides, Pachycephala pectoralis . 

ornata, Holostaspella 

ornatus, Holostaspis 

ornata, Pachycephala pectoralis 

Ornitholestes 

Ostroumovia 

ottomeyeri, Pachycephala pectoralis 



51, 



197 
226 

195 

460 

28 

476 

481 

284 

46, 48 

4, 48 

493 

197 

52, 53 

5i 

204 

60 

483. 506 

• 203 



oudemansii, Macrocheles (Monoplites) 4, 16, 36 
owasianus, Carapus .... 278 



Pachycephala .... 

pallida, Zosterops .... 
palustris, Macrocheles (Monoplites) . 
pannosus, Macrocheles (Nothrholaspis) 
Parholaspis ..... 
parmulatus, Macrocheles (Nothrholaspis) 
parvibrachium, Carapus (Onuxodon) 
parvipinnus, Carapus 
pectoralis, Pachycephala. 
pectoralis, Pachycephala pectoralis 
Pelagohydra .... 
Pelagohydrinae 
pelengensis, Pachycephala pectoralis 



131-222 
• 3i4 



penicilliger, Macrocheles 
penola, Myriothela 
peregrinus, Hypolytus 
phrygia, Myriothela 
pindae, Carapus 
pisentii, Macrocheles 
planctonis, Sagitta 
plumipes, Macrocheles 
plumiventris, Macrocheles 
Podocoryne . 
poeyi, Evoxymetopon 
Porpita. 

primarius, Acaulis . 
producta, Stauridiosarsia 
prolifer, Hybocodon 
prolifera, Sarsia 
Protoholaspinae 
Protoholaspis 
Ptilocodiidae 
Ptilocodium 
punctoscutatus, Macrocheles 
pusilla, Coryne 



12, 27, 



41 
41 

9 

41 

. 284 

. 279 

131-222 
200 

• 508 

• 522 

• 203 
4 1 , 53 

• 467 
460, 506 

4 8 7, 523 

. 277 

11, 13, 14 

437-540 

36, 38, 53 

, 36- 3 8, 41, 53 

. 476 

.• 101 

• 519 
466,481,509,523 

4 6 3. 498, 510 
491, 496, 505 
493. 5i9 
9 
9 

• 525 
474. 513 

18, 19, 53 
498, 525 



queenslandica, Pachycephala pectoralis . 200 



radiatum, Cladonema 

ramosum, Eudendrium 

Rathkea 

reedi, Carapus 

regalis, Tubularia 

rendahli, Carapus 

rendovae, Zosterops 

repens, Ptilocodium 

Rhea . 

Rhizogeton . 

Rhizorhagium 

Riparia 

riparia, Riparia 

Rosalinda 

roseum, Rhizorhagium 

rothamstedensis, Macrocheles 

rufa, Formica 

ryniensis, Asyncoryne 



460, 



511, 519, 525 

• 476 
. 481 
. 287 

• 496 
. 294 

• 316 
5i3, 525 

.63,64 

• 476 

• 476 

• 36 

• 36 

• 5i3 

• 476 
n.15,53 

13 
459, 5°9. 524 



493, 



136 



sagamianus, Encheliophis (Jordanicus) 
Sagitta ..... 

salomonis, Pachycephala 
sanfordi, Pachycephala pectoralis 
Sarsia ...... 

savala, Lepturacanthus . 

scaber, Trox, .... 

Scarabaeus ..... 

schlegelii, Pachycephala . 

sculpta, Holostaspis (Holostaspella) 

semiflava, Zosterops semiflava 

semipunctatus, Scarabaeus 

senegalensis, Zosterops . 

sharpei, Pachycephala pectoralis 

simonyi, Benthodesmus 

simplex, Boreohydra . . 507, 

singularis, Tricyclusa 

Solanderiidae .... 

soror, Pachycephala 

Speirops ..... 

Sphaerocerus .... 

spinicauda, Pachycephala pectoralis 
squamata, Clava ..... 

Stauridiosarsia . . 460, 462, 498, 

Staurocoryne . ..... 

stercorarius, Gamasus .... 

stercorarius, Geotrupes 
strangulata, Dipurena 

Struthio 

subbadius, Holostaspis . 
subbadius, Macrocheles . 11, 19 

submotus, Macrocheles . 11, 12 

superbus, Macrocheles . . 9, 

superciliaris, Bougainvillia 



539 

. 301 

437-450 
207 



197 

5i9 
119 

28 

13 

-222 

5i 
403 

13 
3i4 
202 



5i9 
462 



522 

523 

• 526 

136-222 

309-433 

53 

199 

476 

5io 

462 

16 



493, 



498 

63 
19 

. 34. 36, 53 
■ 28, 29, 53 
12, 38, 39 

. 481 



tabarensis, Pachycephala pectoralis . 

taeniatus, Evoxymetopon 

tardior, Macrocheles (Monoplites) 

tardus, Gamasus .... 

tardus, Macrocheles . . 12 

tenuipes, Holaspulus 

Tentoriceps ..... 

tenuis, Benthodesmus 

terreus, Holostaspis 

teysmanni, Pachycephala pectoralis 

Thyrsitina ..... 

tiarella, Halocordyle 

tidorensis, Pachycephala pectoralis . 

torquata, Pachycephala pectoralis 

Trichiuridae 

Trichiurinae 



30, 



• 203 

99 

41 

13, 30 

3i, 53 

9 

no 



Trichiurus 

Tricyclusa 

Tricyclusidae 

Tricyclusoidea 

tridentinus, Macrocheles 

Tringa . 

Trox 

Tubularia 

Tubulariidae 



462, 482 



• 48 
196 
121 

510, 524 

• 197 
. 198 

73-i3o 
112 

• 113 
489, 508 
453-534 

• 522 

30 

. 64 

28 

470, 481, 496 

453-534 



54° 








INDEX 




Tubularoidea 






520 


virens, Zosterops .... 


• 314 


tubulosa, Sarsia 


493 


498 


505 


viridipectus, Pachycephala schlegelii 


• 205 


Turritopsis .... 






476 


vitiensis, Pachycephala pectoralis 


204 


typica, Gotoea 






493 


voltzkowi, Zosterops maderaspatana 


• 403 


urceolus, Branchiocerianthus . 




472 


522 


whitneyi, Pachycephala pectoralis . 


• 199 


utupuae, Pachycephala pectoralis 






204 


williami, Rosalinda 


• 513 


vagabundus, Macrocheles 






5i 


xanthoprocta, Pachycephala pectoralis 


201 


vanikorensis, Pachycephala pectoralis 






202 


xantusi, Lepidopus 


95 


Veigaia .... 








7 






velella, Velella 








519 


Zanclea . . . 474, 499, 


512, 519 


vermicularis, Encheliophis 








295 


Zancleidae ..... 


• 526 


veterrimus, Macrocheles . 








16 


zetesios, Sagitta .... 


437, 443 


violetae, Pachycephala pectoralis 








199 


Zosterops ..... 


309-433 





6 APR 195S BRITISH MITES 

OF THE SUBFAMILY 
MACROCHELINAE TRAGARDH 
(GAMASINA-MACROCHELIDAE) 



G. OWEN EVANS 

AND 

E. BROWNING 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 4 No. i 

LONDON : 1956 



BRITISH MITES OF THE 

SUBFAMILY MACROCHELINAE TRAGARDH 

(GAMASINA-MACROCHELIDAE) 



BY 

G. OWEN EVANS 

AND 

E. BROWNING 



Pp. 1-55 ; Pis. 1-4 ; 85 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 4 No. 1 

LONDON : 1956 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be compiled 
within one calendar year. 

This paper is Vol. 4, No. 1 of the Zoological series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued March, 1956. Price Seventeen Shillings and Sixpence. 



BRITISH MITES OF THE 

SUBFAMILY MACROCHELINAE TRAGARDH 

(GAMASINA-MACROCHELIDAE) 

By G. OWEN EVANS and E. BROWNING 



CONTENTS 

Introduction ..... 
External Morphology 
Classification ..... 
Genus Macrocheles Latr. 

Macrocheles muscaedomesticae (Scopoli 

Macrocheles pisentii (Berl.) . 

Macrocheles rothamstedensis sp. no v. 

Macrocheles glaber (Miiller) . 

Macrocheles punctoscutatus sp. no v. 

Macrocheles subbadius (Berl.) 

Macrocheles insignitus Berl. 

Macrocheles merdarius (Berl.) 

Macrocheles montanus (Willmann) 

Macrocheles carinatus (C. L. Koch) 

Macrocheles penicilliger (Berl.) 

Macrocheles submotus Falconer 

Macrocheles tardus (C. L. Koch) 

Macrocheles decoloratus (C. L. Koch) 

Macrocheles matrius Hull 

Macrocheles plumiventris Hull 

Macrocheles superbus Hull . 

Species incertae sedis . 
Genus Geholaspis Berl. 5. lat. 

Geholaspis (Geholaspis) longispinosus (Kramer) 

Geholaspis (Longicheles) longulus (Berl.) 

Geholaspis (Longicheles) mandibularis (Berl 
Genus Macrholaspis Oudemans 

Macrholaspis opacus (C. L. Koch) 

Macrholaspis dentatus sp. no v. 
Genus Holosiaspella Berl. 

Holostaspella ovnata (Berl. 
Summary 

Acknowledgments 
References 



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SYNOPSIS 
The classification of the Macrochelidae is discussed, with particular reference to the British 
species, and three new species are described and figured. 

zool. 4, 1. 1 



4 BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 

INTRODUCTION 

The recent lists of the genera of the Macrochelidae follow the classification proposed 
by Berlese (1918), the only comprehensive work on the family. Vitzthum (1941) 
recognizes eight genera and eleven subgenera to which must be added the genera 
Neoholaspis Turk, 1948 (? syn : Macrocheles Latr. s. str.) and Andrholaspis Turk, 1948, 
and the subgenera Cyrtocheles Valle, 1953 and Longicheles Valle, 1953 of Geholaspis 
Berl. s. lat. 

The British Macrochelidae have been investigated by Hull (1918 and 1925), 
Falconer (1923 and 1924) and Turk (1946). In 1918, Hull keyed ten species of 
Macrocheles, of which four were considered to be new. Later, in 1925, he described ten 
more new species and proposed a new name (Macrocheles (Monoplites) oudemansii) 
for Macrocheles marginatus Oudemans, 1901 nee Hermann, 1804. He used the chaeto- 
taxy of the dorsal shield and the form of the ventral shields as his chief taxonomic 
criteria and although these characters have been proved subsequently to be useful 
key characters, his descriptions and figures are so inadequate (and often inaccurate) 
that a number of the species cannot be recognised with certainty. 

Falconer's contribution to the study of the family consists of descriptions and 
figures of two new species and the erection of a new subgenus (Dissoloncha) of 
Macrocheles. 

Turk (1946) has described one new species, Coprholaspis anglicus, from under wet 
wood at Reskadinnick, Cornwall, and has keyed British species of the genus Coprho- 
laspis Berl. 

The object of the present work is to redescribe and figure the known British species 
of the Macrochelinae. We have followed Sellnick's interpretation of the species of 
C. L. Koch and Scopoli (Sellnick, 1931 and 1940). 

EXTERNAL MORPHOLOGY 

The following account of the external morphology of the British species of the 
Macrochelinae refers to the adult stages only and will serve as a general introduction 
to the characters of taxonomic importance used in the keys to species. 

Dorsal shield : The dorsal shield in all British species is entire and covers practically 
the whole of the dorsum of the mite. The ornamentation of the shield shows con- 
siderable variety in form, for example, the shield may be faintly reticulated as in 
Macrocheles glaber (Muller) , strongly reticulated and punctate as in Macrocheles tardus 
(C. L. Koch) or densely covered with minute tubercles as in Geholaspis (Longicheles) 
mandibularis (Berl.). Its lateral margin may be smooth or serrated. 

The number of setae on the shield is remarkably constant, twenty-eight pairs in the 
female, except in three species (Macrocheles montana (Willm.), Macrholaspis opacus 
(C. L. Koch) and Macrholaspis dentatus sp.n.) . The chaetotactic pattern and structure 
of the setae (whether simple, serrated or plumose) afford valuable taxonomic criteria. 
Sellnick (1942) and Valle (1955) have already used the chaetotaxy of the shield in 
their works on Macrocheles and Geholaspis s. lat., respectively. The nomenclature 
used for the dorsal chaetotaxy in the present work is given in Text-fig. 5. The twenty- 



BRITISH MITES — SUBFAMILY M ACROCHELI N AE TRAGARDH 5 

eight pairs of setae of the female are divided into four longitudinal rows ; a dorsal 
series (D) of eight pairs, a median series (M) of four pairs, a lateral series (L) of six 
pairs and a marginal series (Mg) of ten pairs. This division is purely artificial and is 
not based on the post-embryonic developmental sequence. In the males, two groups 







Figs. 1-4. The chaetotaxy of the ventri-anal shield in females of Macrholaspis Oude- 
mans (Fig. 1), Macrocheles Latr. (Fig. 2), Holostaspella Berl. (Fig. 3), and Geholaspis 
Berl. (Fig. 4), a, anterior pre-anal seta ; b, median pre-anal seta ; c, posterior pre-anal 
seta ; d, antero-lateral pre-anal seta ; e, postero-lateral pre-anal seta. 



can be recognized, namely, those which have the same chaetotactic pattern as the 
female and those which have a greater number of setae on the shield. In the latter, 
setae are added to the shield through its extension laterally to incorporate a number 
of the setae (the extra-marginal setae) normally situated on the lateral interscutal 
membrane. 

The anterior margin of the shield in the genera Macrocheles, Macrholaspis and 
Geholaspis is gently rounded with the vertical setae (Di) situated medially at a short 



6 BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 




Fig. 5. Chaetotaxy of the dorsal shield in Macrocheles muscaedomesticae (Scopoli), 
female. D1-D8, dorsal series ; M1-M4, median series ; L1-L6, lateral series ; 
Mgi-Mgio, marginal series. 



BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAgARDH 7 

distance from the margin. In Holostaspella, however, the vertical setae are no longer on 
the summit of the shield but on an outgrowth from it (Text-fig. 82). 

In addition to the setae, the shield is provided with a number of pore-like structures. 
There are normally twenty-two pairs of these " pores ". 

Tritosternutn. This structure is well-developed throughout the group and consists 
of a rectangular base, longer than wide, and a pair of long, pilose lacinae. 

Ventral shields. In the female, the ventral shields comprise the sternal, metaster- 
nals, genital, ventri-anal and metapodals. 

The sternal shield is strongly sclerotized and carries three pairs of setae (hi, h2 
and I13) and two pairs of pores. The ornamentation of the shield is usually character- 
istic of a species or a group of species. Berlese (1918) used the ornamentation of the 
shield as a major character for subdividing the subgenus Coprholaspis and introduced 
a system of nomenclature for the lines and punctate areas forming the basic pattern 
(Text-fig. 2). The writers have referred, in the main, to a photograph of the sternal 
shield instead of attempting a description of the ornamentation. The met asternal 
shields are paired and free, i.e. they are not fused with the sternal or endopodal 
shields. Each bears a seta and usually a " pore ". 

The genital shield lies between coxae IV and carries a pair of genital setae. It is 
invariably reticulated and punctured, and is provided with accessory sclerites as in 
the genus Veigaia Oudms. (Gamasina-Veigaiaidae). The lateral sclerites are usually 
well-developed, but the median sclerite is often only weakly sclerotized. 

The region posterior to coxae IV is largely occupied by a ventri-anal shield bearing 
two (Macrholaspis) , three (Macrocheles) , four (Holostaspella) or five pairs (Geholaspis) 
of pre-anal setae. The nomenclature for these setae is given in Text-figs. 3-6. In 
addition to the pre-anals the shield bears the normal three setae associated with the 
anus, namely, the paired para-anals and the post-anal seta. The surface of the shield 
is usually reticulated and punctured. The interscutal membrane between the genital 
and ventri-anal shields may be provided with sclerotised platelets. The met apodal 
shields are relatively small and inconspicuous. 

In the male, the ventral surface may be covered by a sterniti-genital shield in the 
region of coxae I-IV and a separate ventri-anal shield posterior to coxae IV or, by a 
holoventral shield, i.e. a fused sterniti-genital and ventri-anal shield. The genital 
orifice is prae-sternal. 

Stigmata, peritr ernes and peritrematal shields. The stigmata, one on each side of 
the idiosoma, are situated between coxae III and IV. The peritremes are well- 
developed and extend beyond the level of coxae I. Each forms a U-shaped loop in the 
region of the stigma (Text-fig. 6). The peritrematal shield is fused anteriorly with 
the dorsal shield, but is free in its proximal half. 

Gnathosoma. The gnathosoma is typical of that found in the free-living Gamasina. 
Ventrally, it is provided with four pairs of setae and a distinct ventral or capitular 
groove (Text-fig. 7). The latter has five to seven transverse rows of denticles. The 
corniculi are well-developed and the internal malae long and pilose. The salivary 
styli are also prominent. The five free segments of the pedipalp bear simple, rod-like 
or spatulate setae. The chaetotactic formula for the trochanter, femur and genu is 



8 BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 




Figs. 6-9. Macrocheles muscaedomesticae (Scopoli), female. Fig. 6, lateral view. 
Fig. 7, venter of gnathosoma. Fig. 8, tectum. Fig. 9, chelicera. 

Abbreviations : c.s., capitular seta ; d.sh., dorsal shield ; ext. mg., extra-marginal series 
ext.p.r.; external posterior rostrals; int.p.r., internal posterior rostrals; per., 
peritreme; per.sh., peritrematal shield; r., rostral seta; St., stigma. 



BRITISH MITES — SUBFAMILY M ACROCHELI N AE TRAGARDH 9 

2-5-6. The specialized seta on the inner basal angle of the palptarsus is three pronged. 
This segment also bears a conspicuous long, upright, rod-like seta distally. 

The tectum is extremely variable in form. In the majority of the species it consists 
of three anteriorly directed processes. These may be free (Text-fig. 8) or each lateral 
process partly fused with the median (Text-fig. 31). The lateral processes may be 
smooth and entire or divided and serrated. The median process is usually strongly 
setose and bifurcate distally. In Macrocheles superbus Hull and Geholaspis s. str. 
the tectum is simply produced into a single anteriorly directed process, variously 
divided distally (Text-figs. 57 and 64) . 

The chelicerae in both sexes are chelate-dentate. The teeth are massive and ridged, 
or conical and smooth. The pilus dentilis, dorsal seta and pore, lyriform pore and 
ventral setae are well-developed. In the male the movable digit is provided with a 
strong spermatophoral process. This may be long and slender or short and inflated 
(Text-figs. 39 and 60). 

Ambulatory appendages. All the legs are six-segmented with the ultimate segment 
incompletely divided into a metatarsus and tarsus. Leg I is without an ambulacral 
apparatus (pulvillus and claws) and terminates in a number of sensory setae. This is 
a characteristic feature of the family. Legs II-IV, however, have well-developed 
lobate pul villi and claws. In the females, spurs are present on the femur and tarsus 
of leg II in the genus Holostaspella only (Text -fig. 85). The femur, genu and tibia 
of Leg II and often one or more segments of leg IV are spurred in the males. 



CLASSIFICATION 

Tragardh (1952) divided the Macrochelidae into three subfamilies as follows : 

" 1. Metasternal shields connected with the sternal shield through a narrow bridge 

Protoholaspinae Tragardh, 1949 
-. Metasternal shields free . . . . . . . . . . . 2. 

2. Peritrematic shields not fused with the exopodal shields Macrochelinae Tragardh, 1949 
-. Peritrematic shields fused with the exopodal shields . . Areolaspinae nov. sub f am." 

The genus Protoholaspis Tragardh 1949, the only member of the Protoholaspinae, 
is not a typical Macrochelid in that leg I is provided with an ambulacrum and the 
chelicera lacks the characteristic ventral seta. Further, Tragardh was unable to see the 
structure of the specialized seta on the inner basal angle of the palptarsus, so that 
the exact systematic position of the genus must remain in doubt pending the re- 
examination of the type. The family should, therefore, be considered to consist of 
the two subfamilies separated in couplet 2 of the above key. 

The majority of the British species belong to the Macrochelinae ; the Areolaspinae 
being represented by Holaspulus tenuipes Berl. and Parholaspis sp., which are intro- 
duced species that have become established in the Aroid House, Royal Botanic 
Gardens, Kew. The following separation of the four British genera of the Macro- 
chelinae is based on Evans (1956) : 



io BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 

Key to genera 
Females 

1. Femur of leg II armed with strong spurs ; vertical setae situated on an outgrowth of 

the dorsal shield ; ventri-anal shield with four pairs of pre-anal setae Holostaspella Berl. 
-. Femur of leg II unarmed ; vertical setae on the summit of the dorsal shield ; ventri- 
anal shield with 2, 3 or 5 pairs of pre-anal setae . . . . . . 2. 

2. Ventri-anal shield with two pairs of pre-anal setae . . . Macrholapis Oudemans 
-. Ventri-anal shield with more than two pairs of pre-anal setae . . . . 3. 

3. Ventri-anal shield with three pairs of pre-anal setae . . . Macrocheles Latreille 
-. Ventri-anal shield with five pairs of pre-anal setae . . . Geholaspis Berl. s. lat. 

Genus MACROCHELES Latreille 

Macrocheles Latreille, P. (1829). In Cuvier, Regne anim., Ed. 2, 4 : 282. 
Coprholaspis Berlese, A. (191 8). Redia, 13 : 146. 
Nothrholaspis Berlese, A. (1918). Redia, 13 : 169. 
Dissoloncha Falconer, W. (1923). Naturalist, Lond. : 151. 
Monoplites Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15 : 215. 

Berlese (1918) divided the genus Macrocheles into four subgenera, namely, Macro- 
cheles s. str., Coprholaspis, Nothrholaspis and Geholaspis. Sellnick (1940) has shown 
conclusively that Coprholaspis is a synonym of Macrocheles s. str. and, within recent 
years, Geholaspis has been given generic status. The remaining two subgenera have 
been separated, chiefly, by the ornamentation of the sternal shield in the female. In 
the majority of the species of Macrocheles s. str., this shield is ornamented with distinct 
lines or punctate lines whereas in Nothrholaspis the ornamentation takes the form 
of a network of ridges (cf. PL 1, fig. 3 and PL 3, fig. 15). This distinction is not as 
definite as supposed by Berlese and, as is often the case with a general character of 
this nature, an intermediate group, containing species which could belong to either 
subgenus, is apparent. In view of this, and until a more comprehensive study can be 
made of both sexes of the known Macrocheles species, especially those in the Berlese 
Collection, the writers feel it advisible to combine the subgenus Nothrholaspis with 
Macrocheles. 

Ecologically, the British species of this genus may be divided into two groups : 
those living in dung and those inhabiting leaf litter or mosses. The former contains a 
number of phoretic species, e.g. M. muscaedomesticae and Macrocheles glaber (Miiller), 
in which the shape and chaetotaxy of the dorsal shield in the male differs considerably 
from that in the female. 

The genus Macrocheles may be defined as follows : Dorsal shield in both sexes 
entire with usually twenty-eight 1 pairs of setae and twenty-two pairs of " pores." 
Sternal shield in the female with three pairs of setae ; metasternal shields free. Genital 
shield with a pair of setae and accessory sclerites. Ventri-anal shield with three pairs 
of pre-anal setae in addition to the three setae normally associated with the anus. 
Male with sterniti-geniti- ventral shield and separate ventri-anal or with holoventral 

1 Macrocheles montanus (Willmann) has twenty-nine to thirty-one pairs of setae on the dorsal shield, 
and in some males the number of setae may be increased through the lateral extension of the shield to 
incorporate a number of the extra-marginal setae. 



BRITISH MITES — SUBFAMILY M ACROCHELI N AE TRAGARDH ii 

shield. Genital orifice prae-sternal. Hypostome with well-developed corniculi and 
salivary styli. Pedipalps with five free segments ; chaetotaxy of trochanter, femur 
and genu being (2-5-6). Specialized seta on the palptarsus with three prongs. 
Chelicerae chelate dentate with a well developed brush of setae ventro-laterally ; 
movable digit in the male with a strong spermatophoral process. Legs I without 
ambulacra ; legs II and often legs IV, spurred in the male. 

Type : Acarus marginatus Hermann, 1804 (= Acarus muscae domesticae Scopoli, 
1772). 

Key to the Females of the British Species of Macrocheles Latr. 

1. Setae L3 to L5 and Mg3 to Mg6 simple ........ 2. 

— . Setae L3 to L5 and Mg3 to Mg6 serrate, plumose or pencillate 1 . . . . 8. 

2. Setae D8 simple or pilose .......... 3. 

-. Setae D8 short, comb-like (Text-fig. n) ; sternal shield weakly ornamented ; ventri- 

anal shield considerably longer than broad ; fixed digit of the chelicera with a row 

of 5 or 6 small teeth distally (Text-fig. 13) . . Macrocheles pisentii (Berl.) 

3. Vertical setae (Di) stout and plumose or long and setiform (Text-fig. 22). Dorsal 

shield more than 600^ in length . . . . . . . . . 4. 

-. Vertical setae short spine-like (Text-fig. 24). Dorsal shield less than 500^ in 

length ............. 7. 

4. Vertical setae short and stout, plumose distally ; setae D4, Li, L2, L6 and Mgi 

simple (Text-fig. 17) ; tibia I approximately equal in length to tarsus (1 : i-o- 

1-05) 5- 

-. Vertical setae long setiform (Text-fig. 15) with inconspicuous serrations distally ; 
sternal shield densely punctured ; setae D4, Li, L2, L6 and Mgi pencillate ; 
tibia I shorter than tarsus I (1 : 1-2-1-44) • Macrocheles rothamstedensis sp. nov. 

5. Sternal shield with deeply incised transverse median and posterior oblique lines 

(PI. 1, fig. 3) ; dorsal setae, except Di, D4 and L2, simple 

Macrocheles glaber (Muller) 
-. Sternal shield punctured, transverse median and posterior oblique lines poorly 

developed or absent ; setae D4 simple . . . . . . . . 6. 

6. Ventral shields densely punctured (PI. 1, fig. 4) ; setae Mg2, Mg7, Mgo and D8 

distinctly serrated (Text-fig. 21) . . Macrocheles punctoscutatus sp. nov. 

-. Sternal shield with fewer large punctures (PI. 1, fig. 5) ; setae Di, D8 and L2 only 

serrate or pencillate ...... Macrocheles subbadius (Berl.) 

7. Sternal shield ornamented with large punctures (PI. 1, fig. 6) 

Macrocheles insignitus Berl. 
-. Sternal shield weakly ornamented with punctate lines (PI. 2, fig. 7) 

Macrocheles merdarius (Berl.) 

8. Dorsal shield with 2Q 2 or more pairs of setae (Text-fig. 27) ; median pre-anal seta 

on or slightly off the line connecting the anterior and posterior pre-anals (PI. 2, 

fig. 8) ........ Macrocheles montanus (Willmann) 

-. Dorsal shield with 28 pairs of setae . . . . . . . . . 9. 

9. Vertical setae situated in close proximity to each other, so that their bases are more 

or less contiguous (Text-fig. 5) . . . . . . . . .10. 

-. Vertical setae further apart, their bases distinctly separated, i.e. at least the diameter 

of the setal base apart ........... 12. 

10. Lateral and marginal setae slender, plumose only in their distal third (Text-fig. 5) ; 

sternal shield with distinct lines (PI. 1, fig. 1) ; lateral processes of the tectum free 
(Text-fig. 8) ...... Macrocheles muscaedomesticae (Scopoli) 



1 Except in Macrocheles submotus Falconer in which Mg4 is simple and Mg5 and 6 pilose. 

2 This species is characterised by having nine pairs of setae in the D series. 



12 BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 

-. Lateral and marginal setae strongly plumose in their distal two-thirds ; lateral 

processes of the tectum partially fused . . . . . . . u. 

ii. With three pairs of platelets between ventri-anal and genital shields (PI. 2, fig. 9) . 

Macrocheles carinatus (C. L.Koch) 
-. Without platelets between ventri-anal and genital shields ..... 

Macrocheles penicilliger (Berl.) 

12. Setae D2, Mg2 and Mg4 simple (Text-fig. 36) . . Macrocheles submotus Falconer 
-. Setae D2, Mg2 and Mg4 plumose ......... 13. 

13. Setae Mi simple, approximately equal in length to Di and extending beyond the 

bases of D2 by about one-half their length (Text-fig. 42) 

Macrocheles tardus (C. L. Koch) 
-. Setae Mi simple or plumose and considerably shorter in length than Di . . 14. 

14. Dorsal shield less than 950^ in length ........ 15. 

Dorsal shield more than 1 100^ in length . . . . . . . .16. 

15. Setae Mgio, approximately equal in length to D8, are about half the length of Mg9 

(Text-fig. 44) ; external margin of the lateral processes of the tectum smooth 
anterior to the base of the median process (Text-fig. 45) ; punctate areas on the 
sternal shield inconspicuous .... Macrocheles decoloratus (C. L. Koch) 

-. Setae Mgio equal in length to Mg9 (Text-fig. 47) ; external margin of the lateral 
processes of the tectum serrate well beyond the base of the median process (Text- 
fig. 48) ; punctate areas on the sternal shield large, conspicuous 

Macrocheles matrius Hull 

16. Setae Mi plumose and lying in line with setae Di (Text-fig. 50) ; lateral margins of 

the dorsal shield coarsely and unevenly serrated (Text-fig. 51) ; ventri-anal shield 
considerably broader than long and characteristically shaped (PI. 3, fig. 15) 

Macrocheles plumiventris Hull 
-. Setae Mi simple and lying in line with D2 (Text-fig. 56) ; lateral margins of the dorsal 
shield minutely and evenly serrated (Text-fig. 55) ; ventri-anal shield considerably 
longer than broad ....... Macrocheles superbus Hull 

Macrocheles muscaedomesticae (Scopoli) Sellnick. 

Acarus muscae domesticae Scopoli, J. A. (1772). Annus. V. Hist. Nat. : n. 125, 157. 

Acarus marginatus Hermann, J. F. (1804). Mem. Apt. : 76, figs. 

Macrocheles muscae domesticae, Sellnick, M. (1940). Goteborg. Vetensk. Samh. Handl. (5) 6B, 

No. 14 : 78, figs. 
Macrocheles muscaedomesticae, Pereira, C. & de Castro, M. P. (1945). Arq. Inst. Biol. S. Paulo 

16 : 163, figs. 

Female. Dorsal shield reticulated and bearing twenty-eight pairs of setae and 
twenty- two pairs of " pores " (Text-fig. 5). Vertical setae plumose in their distal third 
and lying in close proximity to each other. Setae D5-D7, Mi, M3 and M4 simple. 
The remainder of the setae on the dorsal shield are pilose in their distal third. The 
distribution of the setae and pores is shown in the figure. Extra-marginal setae are 
simple (Text-fig. 6). 

The sternal shield is characteristically ornamented with punctures and ridges 
(PL 1, fig. 1.) All the sternal setae are simple. The metasternal shields are free and 
each carries a simple seta. Genital shield, truncate posteriorly, bears a simple pair of 
setae and is ornamented with punctate lines. The ventri-anal shield (approx. 
368 x 379/^) has a loose network of punctate lines. All the setae on this compound 
shield are simple. The peritrematal shield is free posteriorly ; being separated from 



BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 13 

the exopodal shields by a wide expanse of striated cuticle (Text-fig. 6) . The peritreme 
extends beyond coxa I. 

The gnathosoma is strongly sclerotized and carries four pairs of setae ventrally 
(Text-fig. 7). The ventral groove is provided with six or seven transverse rows of 
denticles. Anteriorly the corniculi and salivary stylets are strongly developed. The 
chaetotaxy of the five free segments of the pedipalp is normal for the free-living 
Gamasina. The specialized seta on the inner basal angle of the palptarsus is three- 
pronged. The lateral processes of the tectum are free ; the median process is bifid 
distally (Text-fig. 8). The chelicerae are massive ; the movable digit being tridentate 
and the fixed bi- or tridentate (Text-fig. 9) . 

Leg I has the tarsus (187/1) longer than the tibia (165^). 

Male. This sex and the immature stages are described and figured by Pereira & 
de Castro (1945). 

Dimensions. Female : length 980-1014/^ ; breadth 570-671/6. Male : length 
750-900/^ ; breadth 450-600/4. 

Habitat and locality. This species is commonly found on Musca domestica 
Linn, and allied species. Pereira & de Castro (1945) state that all instars except the 
larval feed on the eggs of house flies. M. muscaedomesticae is cosmopolitan in 
distribution. 

Macrocheles pisentii (Berl.) 

Gamasus tardus var. Pisentii Berlese, A. (1882). Bull. Soc. ent. Ital. 14: 112, fig. 
Holostaspis Pisentii Berlese, A. (1887). Acari, Myriopoda et Scorpiones, etc., Fasc. 76, N. 1. 

Female. Dorsal shield, minutely punctured, and bearing twenty-eight pairs of 
setae. Vertical setae simple and well separated from each other (Text-fig. 10). Setae 
D8 short and comb-like (Text-fig. n). The remainder of the setae on the dorsal shield 
long and sharply pointed distally. Extra-marginal setae simple. 

Ventrally, the sternal shield (without distinct ornamentation) has three pairs of 
simple setae. Its posterior margin is strongly concave. Met asternal shields small ; 
setae simple. Genital shield with well-developed lateral sclerites, Ventri-anal shield 
(about 285 x 220/1) is considerably longer than broad. All the setae on this shield 
are simple. 

External posterior rostrals are about-one-third the length of the internals. The 
ventral groove has five transverse rows of denticles. The characteristic form of the 
tectum is shown in Text-fig. 12. The fixed digits of the chelicerae are provided with 
one large, grooved tooth and five or more small teeth (Text-fig. 13). The movable 
digit is tridentate ; the middle tooth being large and recurved. 

Leg I has the tarsus (143/O longer than the tibia (132/6). Tarsus II is provided with 
stout spines. 

Male. Unknown. 

Dimensions. Female : length 810-835/^ ; breadth 490-506/6. 

Habitat and locality. This species has been collected from Scarabaeus semi- 
punctatus in Italy. Hull (1918) records it from a nest of Formica rufa at Chopwell, 



i 4 BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 

Durham ; from moss at Ninebanks, Northumberland ; and in dead leaves, Cheshire. 
We have not examined Hull's specimens. The above description and figures 
are based on specimens in the Oudemans Collection, Leiden. 




Figs. 10-13. Macvocheles pisentii (Berl.), female. Fig. 10, dorsal shield ; Fig. n, 
seta D8 enlarged. Fig. 12, tectum. Fig. 13, chelicera. 



1 



BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAgArDH 15 

Macrocheles rothamstedensis sp. nov. 

Female. Dorsal shield, with twenty-eight pairs of setae and twenty- two pairs of 
" pores ", is weakly ornamented with punctate lines forming a polygonal network 
(Text-fig. 14) . The vertical setae are long and slightly pilose distally ; their bases 
are almost contiguous (Text-fig. 15). Setae D4, Li, L2, L6, Mgi and Mgio are 
conspicuously pilose distally, other dorsal setae may show slight pilosity distally. 
All the extra-marginal setae are simple. 




Figs. 14-16. Macrocheles rothamstedensis sp. nov., female. Fig. 14, dorsal shield. 
Fig. 15, setae Di enlarged. Fig. 16, chelicera. 



i6 BRITISH MITES — SUBFAMILY M ACROCHELI N AE TRAGARDH 

The sternal shield is densely covered with punctures which tend to form a pattern 
of punctate lines (PL I, fig. 2). All the sternal setae are simple. The smooth meta- 
sternal setae lie on small narrow shields. The genital shield, with a pair of simple 
setae, is truncate posteriorly and ornamented with a network of punctate lines. The 
lateral sclerites are strongly formed. The ventri-anal shield (approx. 250 x 198/*) is 
longer than broad and ornamented with a loose network of punctate lines. All the 
setae on this shield are simple. The met apodal shields are small and weakly sclerotised. 

The venter of the gnathosoma is minutely punctured. The ventral groove is 
provided with five rows of denticles. Each lateral process of the tectum is free. The 
movable digit of the chelicera is tridentate ; the fixed digit is basically bidentate 
(Text-fig. 16). 

Leg I (approx. 6io/£ in length) with the tibia (iiojlc) evidently shorter than the tarsus 
(143/^). Legs II and III with simple setae ; leg IV with plumose setae on the femur, 
genu, tibia and tarsus. 

Male. Dorsal shield, strongly attenuated posterior to coxae IV, bears thirty 
pairs of setae. The chaetotactic pattern differs from that in the female in the addition 
of two pairs of extra-marginal setae to the shield and the greater distance between 
the verticals. The ornamentation of the shield is similar to that in the female. 

The venter is covered by a punctured holoventral shield bearing nineteen simple 
setae. 

The tectum is basically the same as in the female. The fixed digit of the chelicera 
is tridentate and the movable is provided with two or three small teeth. The sperma- 
tophoral process is about the length of the movable digit. 

Femur, genu and tibia of leg II and the femur and tibia of leg IV are spurred. 
Femur II and trochanter IV have small sclerotized ridges. 

Dimensions. Male : length 590-595^ ; breadth 365-370^. Female ; length 
710-740//- ; breadth 370-375/6. 

Habitat and locality. Seven females and three males from bullock manure, 
Rothamsted Experimental Station, Harpenden, Herts. Holotype female, 1955.10. 
22 . 43 ; allotype male, 1955 . 10 . 22 . 44 and paratypes, 1955 . 10 . 22 . 45-52. 



Macrocheles glaber (Miiller) 

Holostaspis glabra Miiller, J. (i860). K. K. mahr. schles. Ges. Brunn : 178, figs. 

Gamasus stercorarius Kramer, P. (1876). Arch. Naturgesch. 42 : 95, rig. 

Holostaspis badius, Berlese, A. (1889). Acari, Myviopoda etc., fasc. 52, N. 3. 

Macrocheles marginatus var. littoralis Halbert, J. N. (1915). Proc. Roy. Irish Acad. 31, 39 ii : 

67, fig- 
Macrocheles (Coprholaspis) glaber Berlese, A. (1921). Redia 14 : 85. 
1 Macrocheles (Monoplites) oudemansii Hull, J. E. (1925). Ann. Mag. nat. Hist. (9) 

15 : 215. (in part) 
Macrocheles veterrimus Sellnick, M. (1940). Goteborg. Vetensk. Samh. Handl. (5) 6b : 80, figs. 
Coprholaspis anglicus Turk, F. A. (1946). Ann. Mag. nat. Hist. (11) 12 : 791, figs. syn. nov. 

1 Hull proposed this name for Macrocheles marginatus Oudemans, 1901 nee Herman, 1804. This 
" species," however, is a complex of at least two distinct species : the tritonymph being M. glaber and 
the adult, M. plumiventris. 



BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 17 




18 



Figs. 17-20. Macrocheles glaber (Miiller), female. Fig. 17, dorsal shield. Fig. 1! 
setae Di enlarged. Fig. 19, tectum. Fig. 20, chelicera. 



zool. 4, 1. 



18 BRITISH MITES — SUBFAMILY M ACROCHELI N AE TRAGARDH 

Female. The dorsal shield is weakly reticulated and punctured, and carries 
twenty-eight pairs of setae and twenty-two pairs of " pores" (Text-fig. 17). The 
vertical setae, lying in close proximity to each other, are plumose in their distal half. 
Setae D8 are finely pilose, and D4 and L2 spiculate distally. The remainder of the 
setae on the shield are smooth and sharply pointed. The extra-marginal setae are 
simple and curved. 

The tritosternum is normal. The sternal shield is characteristically ornamented 
with lines and punctate areas (PL 1, fig. 3). The transverse anterior, arcuate, trans- 
verse median and oblique lines are well defined. All the sternal setae are simple. The 
metasternal shields are small and the setae simple. The genital shield is strongly 
ornamented and provided with well-developed accessory sclerites. The ventri-anal 
shield (approx. 264-275/4 long x 265-295/4 wide) is ornamented with concentric 
punctate lines. The median pre-anal setae are situated well outside the connecting 
line between the anterior and posterior pre-anals. All the setae on the ventri-anal 
are simple. The met apodal shields are irregular in outline. The peritreme and peri- 
trematal shield are normal for the genus. 

The gnathosoma bears four pairs of setae ventrally. The internal posterior rostrals 
are about four times the length of the externals. The ventral groove has six transverse 
rows of denticles. The pedipalps are normal. The tectum is produced into three distinct 
processes ; the median being divided distally (Text-fig. 19). The chelicerae are strongly 
developed. The fixed digit is bidentate with the proximal tooth large and ridged 
(Text-fig. 20). The pilus dentilis is relatively short and stout. The movable digit is 
trident ate. The dorsal seta is comb-like distally. 

Leg I (585/*) has plumed setae on the dorsal surface of the femur and genu. The 
tibia and tarsus are approximately equal in length (about 115/4). Legs II (510/4), 
III (460/*) and IV (760/4) have a few plumose setae on the femur and tarsus. 

Male. This sex is figured by Berlese (1889). 

Dimensions. Female : length 850-855/4 ; breadth 540-560/4. 

Habitat and locality. This is one of the commonest species of mites found on 
" dor " beetles in Europe, especially on Geotrupes stercorarius Linn. It is also com- 
monly encountered on Muscid flies. 



Macrocheles punctoscutatus sp. nov. 

Female. Dorsal shield densely covered with minute punctures and bearing 
twenty-eight pairs of setae and twenty-two pairs of " pores " (Text-fig. 21). Vertical 
setae coarsely plumose, well separated (Text-fig. 22). The majority of dorsal setae 
smooth and sharply pointed apically ; setae D8, Mg2, Mg8-io distinctly pilose. 
Extra-marginal setae simple. 

Tritosternum normal, lacinae long and pilose. Sternal shield heavily ornamented, 
punctate areas conspicuous (PL 1, fig. 4). All sternal setae simple. Metasternal 
shields small, metasternal setae simple. Genital shield strongly ornamented, genital 
setae simple. Ventri-anal shield (320/4 x 355/4) broader than long with its anterior 
margin lying in close proximity to the genital shield. The ornamentation of the 



BRITISH MITES — SUBFAMILY MACROCHELINAE TRAGARDH 



19 



shield consists of distinct lines forming a loose network, and numerous small punctures. 
The pre-anals, para-anals and post-anal seta are simple. The metapodals are small 
and elongate ; their long axis lying more or less parallel with that of the longer axis 
of the body. 




Figs. 21-22. Macvocheles punctoscutatus sp. nov., female. 
Fig. 22, setae Di and Mi enlarged. 



Fig. 21, dorsal shield, 



The external posterior rostral setae are about one-half the length of the internals. 
The ventral groove has five transverse rows of denticles. The dentition of the 
chelicerae and the structure of the tectum are similar to that in M. glaber. 

Leg I (605^) has the tibia and tarsus of about equal length. 

Dimensions. Length 880/^ ; breadth 605/^. 

Habitat and locality. A single female (holotype 1955 .10.22. 70) from a mole's 
nest at Churcham, Gloucestershire (Coll. R. S. George). 



Macrocheles subbadius (Berl.) 1 
Holostaspis subbadius Berlese, A. (1904). Redia 1 : 264. 

Female. Dorsal shield bears twenty-eight pairs of setae and twenty- two pairs 
of " pores " (Text-fig. 23). The vertical setae are plumose distally. Setae D8 and, 

1 This species is figured by Berlese (1889) under Holostaspis marginatus (Herm.) Berl. " forma inter- 
media inter badium et adultum (marginatum) foem " in Acari Myriopoda, etc., fasc. 52, N. 6. 



20 BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 

in the majority of the specimens examined, setae L2 are pilose. The extra-marginal 
setae are simple. 

The sternal shield is ornamented with distinct punctures arranged as in PI. 1, fig. 5. 
The sternal, metasternal and genital setae are simple. The ventri-anal shield (190-220 
X ic;0/£) is ornamented with punctate lines ; the nine setae on this shield are simple. 
The met apodal shields are weakly sclerotized. 




Fig. 23. Macrocheles subbadius (Berl.), female. Dorsal shield. 



The lateral processes of the tectum are free and the form of the chelicerae is 
essentially the same as in M. merdarius. 

Leg I has the tibia (82/*) shorter than the tarsus (iio/j). The spines on tarsus II 
are short and stout. 

Male. Unknown. 

Dimensions. Length 610-625^ '> breadth 370-400/^. 



BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 21 

Habitat and locality. Berlese (1889) records this species from manure in 
Italy. The writers have examined specimens from farmyard manure, Evesham, 
Worcestershire. 



Macrocheles insignitus Berl. 

Macrocheles (Coprholaspis) insignitus Berlese, A. (1918). Redia 13 : 158. 

Female. Dorsal shield with twenty-eight pairs of setae and twenty-two pairs of 
" pores ". The surface of the shield is covered with minute punctures which form 
a polygonal network. All the setae on the shield and the interscutal membrane are 
simple. The verticals are short spine-like. 

Sternal shield characteristically ornamented with punctures as in PL 1, fig. 6. All 
the sternal setae are simple. The metasternals are small and flank the anterior part 
of the ornamented genital shield. The met asternal and genital setae are simple. The 
ventri-anal shield is about as broad as long (156 x 152/4) and ornamented with 
punctate lines. The setae on this compound shield are all simple. The metapodals 
are extremely weakly sclerotised. 

The gnathosoma is typical for the genus. The lateral processes of the tectum are 
free and fish-tail like ; the median process is strongly bifurcate distally. The form 
of the chelicerae is similar to that in M. merdarius. 

Leg I (approx. 350/4 in length) has the tibia (55/*) considerably shorter than the 
tarsus (72/^). 

The male is unknown. 

Dimensions. Length 445/4 ; breadth 275/4. 

Habitat and locality. This species is previously known by a single female from 
the type locality " Longny, Orne in Gallia" (Berlese, 1918). The writers have 
examined a single female of this species from " a hot bed ", Austrey, Warwickshire 
(undetermined in the Michael Collection). 



Macrocheles merdarius (Berl.) 

Holostaspis merdarius Berlese, A. (1889). Acari, Myriopoda et Scorpiones etc., Fasc. 52, N. 1, fig. 
Macrocheles merdarius, Sellnick, M. (1940). Goteborg. Vetensk. Samh. Handl. (5) 6b : 86, figs. 

Female. Dorsal shield, ornamented with reticulations, is provided with twenty- 
eight pairs of simple setae and twenty-two pairs of pores (Text-fig. 24). The vertical 
setae are sub-spinose. The extra-marginal setae are also simple. 

The sternal shield is lightly ornamented with punctate lines and the three pairs of 
sternal setae are simple (PI. 2, fig. 7). The metasternal shields are minute. Both the 
met asternal and genital setae are simple. The truncated anterior margin of the 
ventri-anal shield lies in close proximity to the posterior margin of the genital. The 
ventri-anal (about 150 x 126/1) is ornamented with four or five transverse lines. All 
the setae on this shield are simple. The metapodals are small and weakly sclerotized. 



22 BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 

Ventral groove with five transverse rows of denticles. External posterior rostrals 
about one-third the length of the internals. The tectum has the lateral processes free 
(Text-fig. 25). The median process is deeply bifurcate. The dentition of the digits 
of the chelicerae is shown in Text-fig. 26. 






Figs. 24-26. Macvocheles merdarius (Berl.), female. Fig. 24, dorsal shield. 
Fig. 25. tectum. Fig. 26, chelicera. 



Male. This sex is described and figured by Berlese (1889). 

Dimensions. Female : length 445-490/^ ; breadth 225-280/^ Male : length 
380/4 ; breadth not given by Berlese (1889). 

Habitat and locality. This is one of the commonest species of Macrochelids 
occurring in dung and compost. It has been recorded from a number of localities in 



BRITISH MITES — SUBFAMILY M ACROCHELI N AE TRAGARDH 



23 



Europe (Franz, 1954). In Britain it is recorded from bullock dung at Rothamsted 
Experimental Station, Harpenden, Herts. (Hyatt, 1956). 

Macrocheles montanus (Willmann) 
Nothrholaspis montana Willmann, C. (1951). Bonner Zool. Beitr. 2 : 158, figs. 

Female. Dorsal shield with twenty-nine pairs of setae and twenty- two pairs of 
" pores " (Text-fig. 27). The surface of the shield is conspicuously punctated and 




Fig. 27. Macrocheles montanus (Willmann), female. Dorsal shield. 



24 BRITISH MITES — SUBFAMILY MACROCHELIN AE TRAGARDH 

reticulated. Dorsal series (D) comprising nine pairs of setae. Vertical setae strongly 
plumose with their bases well separated. Setae Mi, D5-D8, M3 and M4 and the 
additional setae of the D series are simple. The remainder of the dorsal setae are 
plumose. Extra-marginal setae are simple. 

The sternal shield is covered with punctures which are larger in the posterior third 
of the shield (PI. 2, fig. 8). The first pair of sternal setae are plumose in their distal 
third ; the second and third pairs are smooth or slightly plumose distally. The form 
of the metasternal and genital shield is shown in the figure. Ventri-anal shield (357 x 




Fig. 28. Macrocheles montanus (Willmann), female. Chelicera. 



340/4) is ornamented with a network of punctate lines. The median pre-anal seta lies 
almost on the line connecting the anterior and posterior pre-anals. The pre-anal and 
para-anal setae are simple, the post-anal seta is strongly plumose distally. There are 
three pairs of platelets between the genital and ventri-anal shields. The metapodal 
shields are weakly sclerotized. 

Venter of the gnathosoma normal for the genus. Tectum essentially the same as in 
M. submotus (Text-fig. 37). Both digits of the chelicerae tridentate (Text-fig. 28). 

Leg I with the tarsus (175/4) longer than the tibia (154/4). 

The male is unknown. 

Dimensions. Length 1,050-1,140/4 ; breadth 660/4. 

Habitat and locality. This species is previosuly known from Austria only 



BRITISH MITES — SUBFAMILY M ACROCHELI N AE TRAGARDH 25 

(Willmann, 1951 and Franz, 1954). We have examined a single female collected 
under " old wet oak bark " in Marley Wood, Wytham, Berkshire (Coll. E. W. Fager). 
M. montanus is the only British species in the genus which has more than the normal 
compliment of setae on the dorsal shield. In Willmann's figure of the type specimen 
there are thirty-one pairs of setae on the shield ; additional plumose setae being 
present near setae L7 and L9 as well as the simple pair in the D series. In other 
characters the British specimen agrees with the type. 



Macrocheles carinatus (C. L. Koch). 

Gamasus carinatus Koch, C. L. 1839. Deutsch. Crust. Myr. Arach. fasc. 24, t. 16. 
Macrocheles hypochthonius Oudemans, A. C. 191 3. Ent. Ber. Amst. 4 : 6. 
Macrocheles hypochthonius Oudemans, A. C. 1914. Arch. Naturgesch. 79a, Hft. 8 : 175, figs. 
Nothrholaspis hulli Falconer, W. 1923. Naturalist Lond. : 153, figs. syn. nov. 
Nothrholaspis carinata Sellnick, M. 1931. SB. Akad. Wiss, Wien 140 : 766, figs. 

Female. The dorsal shield is strongly reticulated and punctured, and its lateral 
margins serrated (Text-fig. 29) . The vertical setae (Di) which stand in close proximity 
to each other, are long and strongly plumose in their distal half (Text-fig. 30) . Setae 
Mi are short and may be smooth or plumose. Setae M3, M4, D6, D7, and D8 are 
thin and simple or slightly serrated distally. The remainder of the setae on the 
dorsal shield are strong and plumose. Setae D3 lie in advance of M2. The anterior 
extra-marginal setae are simple and the posterior plumose. 

The sternal shield is ornamented with a network of ridges and punctures ; the latter 
are especially large in the posterior third of the shield (PL 2, fig. 9). The sternal and 
metasternal setae are simple. The posterior margin of the genital shield is strongly 
convex, the genital setae are simple. The ventri-anal shield is sub-circular in outline 
and is strongly ornamented. All the setae on the compound shield are simple. The 
median pre-anals lie considerably closer to the anterior than the posterior pre-anals 
and are situated almost on the connecting line between these setae. The interscutal 
membrane between the genital and ventri-anal shield bears six platelets. The 
met apodal shields are small. The peritreme and peritrematal shield are normal for 
the genus. The interscutal membrane is conspicuously corrugated. 

The external posterior rostral setae are about one-third the length of the internals. 
The ventral groove carries five rows of denticles. The tectum (Text-fig. 31) has the 
lateral lobes partially fused ; the lateral and median lobes are divided distally. The 
chelicerae are strongly developed with the fixed digit tri-dentate (Text-fig. 32). The 
movable digit may be bi- or tri-dentate. The dorsal seta is serrated on one side. 

Leg I, approximately 935/4 in length, has the tibia (120/*) considerably shorter than 
the tarsus (165/4). Legs II, III, IV measure approximately 660//, 605/4, and 1,100/4 
respectively. 

Male. The chaetotaxy of the dorsal shield is basically the same as in the female. 
The sterniti-genital shield is provided with a polygonal network of ridges and strong 
punctures. The ventri-anal shield is also reticulated and punctured and measures 
about 298/4 in length and 310/4 in breadth. All the setae on this shield are simple. 



26 



BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 



The tectum is similar to that in the female. The spermatophoral process on the 
movable digit of the chelicera is short, being about one-half the length of the digit. 

Femur, genu and tibia of leg II only are spurred. 

Dimensions. Male: 880-900/1 in length ; 500-540/^ in breadth. Female: 1,034- 
i,oy8/i in length ; 570-640/t in breadth. 

Habitat and locality. In humus and moss from a number of localities in the 
British Isles. This species has also been recorded from Austria, Germany and the 
Netherlands. 




<^y 





Figs. 29-32. Macvocheles carinatus (C. L. Koch), female. Fig. 29, posterolateral 
margin of the dorsal shield. Fig. 30, anterior region of the dorsal shield. Fig. 31, 
tectum. Fig. 32, chelicera. 



BRITISH MITES — SUBFAMILY M ACROCHELI N AE TRAGARDH 



27 



Macrocheles penicilliger Berl. 

Holostaspis penicilliger Berlese, A. (1904). Redia 1 : 264. 

Holostaspis penicilliger Berlese, A. (1918). Redia 13 : 146, 162. 

Macrocheles penicilliger, Sellnick, M. (1940). Goteborg. Vetensk. Sarnh. Handl. (5) 6b : 82, figs. 

Female. The dorsal shield is strongly reticulated and punctured, and has its 
lateral margins serrated (Text-fig. 33) . Vertical setae, approximately 55//, in length, 
are plumose in their distal two-thirds. Their bases are almost contiguous. Setae Mi, 
M3, M4, D6-D8 are simple ; remainder of dorsal setae plumose. Setae D8 are situated 
considerably anterior to the line connecting setae Mgio. Extra-marginal setae are 
plumose distally. 




Figs. 33-35. Macrocheles penicilliger (Berl.), female. Fig. 33, dorsal shield. 
Fig. 34, tectum. Fig. 35, chelicera. 



28 BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 

The sternal shield is heavily ornamented with ridges and punctures (PI. 2, fig. 10) 
The first pair of sternal setae are strongly plumose and the second and third pair 
slightly plumose distally. The metasternal setae are simple as are the pair of setae 
situated on the punctured genital shield. On the ventri-anal shield (approx. 310 x 
330/^), the pre-anal and para-anal setae are simple and the post-anal seta plumose. 
The median pre-anal lies outside the line connecting the anterior and posterior 
pre-anal. There are no platelets between the ventri-anal and genital shields. The 
metapodals are strongly sclerotized and elongate. 

The external posterior rostrals on the venter of the gnathosoma are about one- 
third the length of the internals. Ventral groove has five transverse rows of denticles. 
The lateral processes of the tectum are partially fused ; distally these processes may 
or may not be divided (Text-fig. 34). The median process is deeply bifurcate. Both 
digits of the chelicerae are bidentate (Text-fig. 35). The pilus dentilis is short and 
thick at its base. The ventral seta is long and strongly pilose. 

Leg I with tibia (approx. 99/^) considerably shorter than the tarsus (approx. 130/*). 

Male. Unknown. 

Dimensions. Female : length 900-930/4 ; breadth 525-550/4. 

Habitat and locality. This species has been found on Trox scaber (Linn.) in 
the nests of owls at Wytham, Oxford (Coll. C. E. Elton) and at Woodford, Essex 
(Coll. A. Hooper). Further records are from decaying leaves, Waterworks Valley, 
Jersey (Coll. G. Owen Evans), Italy (Berlese, 1904), Iceland (Sellnick, 1940) and 
Austria (Franz, 1954). 

Macrocheles submotus Falconer, 

Macrocheles cognatus Falconer, W. (1923). Naturalist, Lond. : 152, fig. (nom. praeocc). 
Macrocheles submotus Falconer, W. (1924). Naturalist, Lond. : 363 (nom. nov. pro. M. cognatus). 
Macrocheles (Nothrholaspis) occidentalis Hull, J. E. (1925). Ann. Mag. nat. Hist. (9) 15 : 213, 

fig. syn. nov. 
? Macrocheles (Nothrholaspis) gloriosus Hull, J. E. (1925). Ann. Mag. nat. Hist. (9) 15 : 214, 

fig. syn. nov. 

Female. Dorsal shield, strongly reticulated and punctured, is provided with 
twenty-eight pairs of setae and twenty- two pairs of pores. Its lateral margins are 
entire. Setae Di plumose and situated about one diameter of their bases apart 
(Text-fig. 36). Setae Mi, M4, D2, D5-D8, Li 1 , Mg2 and Mg4 are simple, and Mg5 and 
Mg6 pilose. The remainder of the setae on the dorsal shield are distinctly plumose. 
The extra-marginal setae are simple or slightly pilose. 

The tritosternum is normal with strongly pilose lacinae. The sternal shield, 
ornamented with strong ridges and large punctures, bears three pairs of simple, 
spine-like setae (PI. 2, fig. 11). The metasternal and genital setae are also simple. 
Ventri-anal shield (approx. 450 x 43 0/1) is triangular in outline and ornamented 
with ridges and punctures. The pre-anal and para-anal setae are simple. The median 
pre-anal lies outside the line connecting the anterior and posterior pre-anals. Three 
pairs of platelets lie between the ventri-anal and genital shields. The interscutal mem- 
brane is closely striated. The peritreme and peritrematal shields are normal. 

1 This seta should be simple in Text-fig. 36. 



BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 



29 




Figs. 36-39. Macrocheles submotus Falconer. Fig. 36, dorsal shield of female. 
Fig. 37, tectum of female. Fig. 38, chelicera of female. Fig. 39, chelicera of male. 



3 o BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 

The external posterior rostrals on the venter of the gnathosoma are about one-half 
the length of the internals. The ventral groove carries five transverse rows of denticles 
The form of the tectum is shown in Text-fig. 37. Both digits of the chelicerae are 
massive (Text-fig. 38). The fixed digit has two strong and two weaker teeth ; the 
movable digit is basically bi-dentate. The dorsal seta is comb-like. 

Leg I (approx. 1,100/* in length) has the tibia (205/*) shorter than the tarsus (220/*). 
Legs II-IV measure respectively about 1,150/*, 1,078/* and 1,700/*. 

Male. The chaetotaxy and ornamentation of the dorsal shield is essentially the 
same as in the female. Ventrally, the truncated sterniti-genital shield is coarsely 
punctured. The five pairs of setae on this compound shield are simple. The ventri- 
anal shield (350 x 340/*) is reticulated and punctured ; there are no platelets between 
the ventri-anal and genital shield. Both digits of the chelicerae are dentate as in 
Text-fig. 39. The spermatophoral process is considerably shorter than the length of 
the digit. 

The femur, genu and tibia of leg II only are spurred. 

Dimensions. Male : length 1,050-1,100/* ; breadth 620-630/*. Female : 1,390- 
1,450/* ; breadth 780-820/*. 

Habitat and locality. This is one of the commonest, and most widely distributed 
Macrochelid found in litter and humus under deciduous and coniferous trees in 
Britain. 

The above description of submotus is based on specimens compared with the type 
in the Falconer Collection at the Liverpool Museum. This species has also appeared 
under the name of Macrocheles tridentinus (Can.) in British and possibly other 
European faunal lists. The original description of tridentinus was based on the male 
only and both description and drawings are insufficient for the certain identity of the 
species. Canestrini gives the length of the male of tridentinus as 860/*. 



Macrocheles tardus (C. L. Koch) 

Gamasus tardus Koch, C. L. 1841. Deutsch. Crust. Myr. Arach. fasc. 39, t. 14. 
Nothrholaspis tardus, Sellnick, M. 1931. S.B. Akad. Wiss. Wien. 140 : 765, figs. 

Female. Dorsal shield, strongly reticulated and punctured, is provided with 
twenty-eight pairs of setae and twenty- two pairs of pores. The vertical setae (Di) 
are plumose and well separated, setae Mi are long, simple and extend beyond the 
bases of D2 by about one-half their length (Text-fig. 40). Setae M3, M4, D6 and D8 
are simple, D7 are finely pectinated whilst the remainder of the setae of the dorsal 
shield are strongly plumose. The marginal setae (Mg) are situated a short distance 
from the lateral margin of the shield (Text-fig. 41). The anterior extra-marginal 
setae are pectinate and the posterior extra-marginals plumose. 

The tritosternum is normal with strongly pilose lacinae. The sternal shield is 
ornamented with a polygonal network of ridges and numerous punctures (PL 2, fig. 
12). All the sternal setae are simple. The metasternal shields are small and the setae 
plumose. The strongly punctured genital shield is convex posteriorly and bears a 
pair of simple setae. The ventri-anal shield, about as broad as long (330 x 300/*) is 



BRITISH MITES — SUBFAMILY M ACROCH ELI N AE TRAGARDH 



3i 



reticulated and punctured. The anterior and posterior pre-anals are simple but the 
median pre-anal may be simple or pectinate. The para-anals are long and simple and 
the post-anal seta short and plumose. The region between the genital and ventri-anal 
shields is usually provided with three pairs of platelets. This number may be reduced 
in some specimens. The metapodal shields are small. The peritreme and peritrematal 
shield are normal. The interscutal membrane is coarsely striated, the striae being 
provided with triangular processes at intervals along their length. 

Ventrally, the gnathosoma has the normal four pairs of setae of which the external 
posterior rostrals are about one-half the length of the internals. The ventral groove 







Figs. 40-43. Macrocheles tardus (C. L. Koch), female. Fig. 40, anterior region of the 
dorsal shield. Fig. 41, postero-lateral margin of dorsal shield. Fig. 42, tectum. 
Fig. 43, chelicera. 



32 BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 

has five rows of denticles. The lateral processes of the tectum are partially fused 
(Text-fig. 42) ; the distal end of these processes being bi- or trifurcate. The stout 
median process is divided ; each arm being entire or divided distally. The fixed digit 
of the chelicera is provided with two strong teeth and between them two or three 
weaker teeth (Text-fig. 43). The pilus dentilis is long and stout. The movable digit 
has two recurved teeth only. The dorsal seta is large and comb-like. 

Leg I (approximately 1,020/t in length) bears plumose setae on the femur, genu and 
tibia. The tibia (165-175/1) is shorter than the tarsus (200-215/*). Leg II (880/1) has 
plumose setae on the trochanter and tarsus as have leg III (820/*) and leg IV 1,485/*). 

Male. The chaetotaxy and ornamentation of the dorsal shield is essentially the 
same as in the female. The sterniti-genital shield, truncated posteriorly, is heavily 
reticulated and punctured. The five pairs of setae on the compound shield are 
simple. The ventri-anal shield (330 x 350/*) is also reticulated and punctured. The 
pre-anal and para-anal setae are usually simple, but may be pectinate. The post- 
anal seta is strongly pilose. The chelicerae are basically the same as in M. cognatus, 
differing only in the position of the large proximal tooth on the fixed digit. The 
spermatophoral process is considerably shorter than the length of the movable digit. 

The femur, genu and tibia of leg II are the only segments which are spurred. 

Dimensions. Male : 1,030-1,100/* in length ; 627-700/* in breadth. Female 
1,215-1,290/* in length ; 730-800/* in breadth. 

Habitat and locality. In vegetable debris near Rydal Water, Westmorland 
and in decaying leaves, Waterworks Valley, Jersey. This species has also been 
recorded from a number of other localities in Europe (Franz, 1954). 



Macrocheles decoloratus (C. L. Koch) 

Gamasus decoloratus Koch, C. L. (1893). Deutsch. Crust. Myr. Arach. fasc. 25, t. 14. 
Macrocheles decoloratus, Oudemans, A. C. (1913). Ent. Ber. Amst. 4 : 5. 
Macrocheles decoloratus, Oudemans, A. C. (1914). Arch. Naturgesch. 79a Hft. 8 : 173. 

Female. Dorsal shield, weakly reticulated but strongly punctured, bears twenty- 
eight pairs of setae and twenty- two pairs of pores (Text-fig. 44). The vertical setae 
(Di) are well separated and strongly plumose. The remainder of the setae on the 
dorsal shield are also plumose. A characteristic feature of the chaetotaxy is the 
relative length of setae Mg9 and 10. The latter is approximately equal in length to 
D8, but only about one-half the length of M.gg. The extra-marginal setae are plumose. 

The tritosternum is normal. The sternal shield (195/* long x 220/* wide) is orna- 
mented with a faint reticulate pattern and large punctures (PI. 3, fig. 13). The punc- 
tate areas are usually conspicuous. The three pairs of sternal setae and the metaster- 
nals are long and simple or slightly pectinate. The genital shield is strongly punctured 
and has a network of punctate lines in its anterior half. The genital setae usually are 
simple. The ventri-anal shield (286-298/* long x 250-286/* wide) is provided with 
a network of punctate lines. The lateral regions of the shield are densely punctured. 
The pre-anal and para-anal setae are long and simple. The post-anal seta is shorter 
and plumose. There are no platelets between the genital and ventri-anal shield. The 



BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 



33 



metapodals are small and irregular in outline. The peritreme and peritrematal shields 
are normal. 

Ventrally, the gnathosoma carries the normal four pairs of setae. The external 
posterior rostrals are about one-third the length of the internals. The capitular 




MglO 



Fig. 44. Macvocheles decoloratus (C. L. Koch), female. Dorsal shield. 



groove is provided with five transverse rows of denticles. The tectum (Text-fig. 45) 
has the three anterior processes separate. The movable digit of the chelicera is 
tridentate and the fixed bi- or tridentate (Text-fig. 46). The dorsal seta is spatulate. 

Leg I (approx. 6yo/i) with plumose setae on femur, genu and tibia. Tarsus I 
(132-140/*) longer than tibia I (121/1). Leg II (680/*). Leg III (638/*) and Leg IV 
(ggojLt) with some plumose setae on trochanter and tarsus. 

Male. Unknown. 

ZOOL. 4, I. 3 



34 



BRITISH MITES — SUBFAMILY MACROCHELIN AE TRAGARDH 



Dimensions. Length 850-880/6 ; breadth 510-520/4. 

Habitat and locality. In cow dung, nr. Canterbury, Kent (Coll. E. Warren, 
1942). This species has also been recorded from Austria, Germany and the Nether- 
lands. 





45 



Figs. 45 — 46. Macrocheles decoloratus (C. L. Koch), female. 

Fig. 46, chelicera. 



Fig. 45, tectum. 



Macrocheles matrius (Hull). 

Nothrholaspis matrius Hull, J. E. (1925). Ann. Mag. nat. Hist. (9) 15 : 212. 

Macrocheles subbadius var. robustulus, Sellnick, M. (1940). Goteborg. Vetensk. Samh. Handl. 

(5) 6b : 86, figs. 
Macrocheles carinatus, Hughes, A. M. (1948). Mites associated with stored food products. 

H.M.S.O. : 126, figs. 

Female. Superficially, the chaetotaxy of the dorsal shield in this species resembles 
that in M. decolor atus. The chief difference between the species lies in the relative 
lengths of setae Mgo, and Mgio which in the present species are approximately equal 
in length (Text-fig. 47). The extra-marginal setae are slightly plumose. 

The sternal shield (187/6 long x 220/6 wide) has well developed lines and a few 
punctures (PL 3, fig. 14). The punctate areas are considerably larger and more 
strongly developed than in M. decolovatus. The sternal and met asternal setae are 
simple or slightly pectinate. The genital shield is ornamented with punctate lines and 
large punctures. The genital setae are plumose distally. The ventri-anal shield 



BRITISH MITES — SUBFAMILY M ACROCHELI N AE TRAGARDH 



35 



(340/6 x 375/^ wide) is characteristically ornamented. The pre-anal and para-anal 
setae are simple. The post-anal seta is short and plumose distally. Metapodals are 
elongate and weakly sclerotized. 

The gnathosoma is essentially the same as in decolor atus except for the structure of 




MglQ 

Fig. 47. Macvocheles matrius Hull, female. Dorsal shield. 



the tectum ; in matrius the external margin of the lateral processes are strongly 
serrulate (Text-fig. 48). 

Male. This sex is adequately described and figured by Hughes (1948). 

Dimensions. Male : length 680-720/t ; breadth not stated by Hughes (1948). 
Female : length 890-910/t ; breadth 540-565/*. 

Habitat and locality. Hull (1925) states that the species is " not uncommon in 
north of England. Abundant in poultry manure, West Allendale ". We have 



36 BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 

examined specimens from the nest of Riparia riparia at Tirley, Gloucestershire 
(Coll. R S. George) and from poultry manure at Houghton, Huntingdonshire 
(Coll. C. Horton Smith). Hughes (1948) records it under the name M. carinatus 
from floor debris and sievings of grain. 




Fig. 48. Macrocheles matrius Hull, female. Tectum. 

M. matrius is undoubtedly conspecific with Macrocheles subbadius var. robustulus 
(Berl.) Sellnick (1940). If the species described and figured by Sellnick is the same as 
robustulus, there is some doubt concerning this, then matrius must be considered a 
synonym. 

Macrocheles plumiventris Hull 

Holostaspis marginatus Berlese, A. (1889). Acari, Myriopoda et Scorpiones, etc., fasc. 52, No. 

4 and 5, figs. 
? Macrocheles gladiator Hull, J. E. (1918). Trans. Nat. Soc. Northumb., n.s. 5 : 71, figs. 
? Macrocheles plumipes Hull, J. E. (1918). torn. cit. : 72, fig. 
Macrocheles (Monoplites) oudemansii Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15 : 215 

(in part). 
Macrocheles plumiventris Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15 : 216, figs. 
Nothrholaspis fimicola Sellnick, M. (1931). S.B. Akad. Wiss. Wien 140 : 765, fig. syn. nov. 

Female. The dorsal shield is strongly reticulated, especially in its posterior 
two- thirds. The lines of the mesh-work are distinctly crenulated (Text-fig. 49). The 
vertical setae (Di) which are well separated, and setae D2-D4, M2, series L and series 
Mg. are strongly plumose in their distal half (Text-fig. 50). Setae Mi are short and 
relatively weakly plumose ; setae D6-D8 and Mg2-3 are considerably finer and less 
plumose than the L and the remainder of the Mg. series. The marginal setae are 
situated on the serrated lateral margin of the dorsal shield (Text-fig. 51). The 
extra-marginal setae are plumose. 

The tritosternum is normal. All the ventral shields are strongly reticulated and 



BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 

I l D1 ^ 



37 





54 






Figs. 49-54. Macrocheles plumiventris Hull, female. Fig. 49, ornamentation of the 
dorsal shield. Fig. 50, anterior region of the dorsal shield. Fig. 51, lateral margin 
of the dorsal shield. Fig. 52, tectum. Fig. 53, chelicera. Fig. 54, dorsal seta on 
chelicera 



3S 



BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 



punctured (PL 3, fig. 15). The sternal shield bears three pairs of sternal setae of 
which hi is strongly plumose and I12-3 smooth and blunt. The met asternal shields 
are free and the setae plumose. The genital shield is provided with accessory sclerites 
and the genital setae are plumose. The ventri-anal shield is considerably broader than 
long (370-460^ x 490-610/4) and characteristically shaped. The three pairs of pre- 
anal setae are plumose, the para-anal setae fine and slightly pilose, and the post-anal 
seta short and strongly plumose. The metapodal shields are well developed. The 
peritreme and peritrematal shield are normal for the genus. 

Ventrally, the gnathosoma bears four pairs of setae. The external posterior 
rostrals are more than one-half the length of the internal posterior rostrals. The 
ventral groove is provided with seven rows of denticles. The pedipalps are normal. 
The tectum has its lateral processes partially fused. The apex of these processes may 
be entire or bifurcated (Text-fig. 52). The median process is strongly bifid distally. 
The fixed digit of the chelicera has five teeth of which the larger are ridged (Text-fig. 
53). The movable digit is provided with three teeth, the proximal being the smallest. 
The dorsal seta has a denticulate margin (Text-fig. 54). The ventral setae are long 
and strongly pilose. 

Leg I, about 1,200/6 in length, has plumose setae on the femur, genu and tibia. 
Trochanter I bears a short conical spur internally. Tibia I measures 220-242/t and 
Tarsus I 210-220/4. Leg II is stout, approximately 1,100/4 in length and richly 
provided with plumose setae as are leg III (approx. 1,030/6) and Leg IV (approx. 
1,650/4.). The pulvilli and claws of Legs II— III are well developed. 

Male. The ornamentation and chaetotaxy of the dorsal shield is basically the 
same as in the female. The venter is covered by a reticulated holoventral shield. 
The fixed digit of the chelicera is provided with four teeth in its distal half. The 
movable digit, in the specimen under study, has a single tooth. Berlese (1889) shows 
four teeth on this digit ! The spermatophoral process is longer than the movable 
digit and is pointed distally. The trochanter, femur, genu, tibia and tarsus of Leg II 
are provided with spurs, also genu III, and trochanter and femur IV. 

Dimensions. Male : 913/4 in length and 605/4 in breadth (a single specimen in 
the Michael Collection). Female : 1,150-1, 480/1 in length and 750-935/4 in breadth. 

Habitat and locality. In manure and compost heaps. Common and widely 
distributed in Europe. 

Sellnick (1931) drew attention to the fact that Holostaspis marginahis Berl. (1889) 
was not Acarus marginatus Hermann (1804) and proposed the name Nothrholaspis 
fimicola for Berlese's species. Hull (1925) had, however, already described this species 
under Macrocheles plumiventris. It is also possible that Hull's Macrocheles gladiator 
and plumipes are referable to this species, but the original descriptions and figures 
are wholly inadequate for their certain identity. 

Macrocheles superbus Hull 
Macrocheles superbus Hull, J. E. (1918). Trans. Nat. Hist. Soc. Northumb., n.s. 5 : 71, figs. 

Female. Dorsal shield minutely punctured and with a faint reticulated pattern 
in its posterior half. The lateral margin of the shield is evenly serrated (Text-fig. 55). 



BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 39 




Figs. 55-62. Macrocheles superbus Hull. Fig. 55, lateral margin of dorsal shield. Fig. 
56, dorsal shield of female. Fig. 57, tectum of female. Fig. 58, chelicera of female. 
Fig. 59, distal end of tectum of male. Fig. 60, chelicera of male. Fig. 61, leg II of 
male. Fig. 62, leg IV of male. 



4 o BRITISH MITES — SUBFAMILY MACROCHELINAE TRAGARDH 

The vertical setae (Di) are stout, plumose and widely separated (Text-fig. 56). Setae 
D4-D8, M3 and M4 are simple and minutely spiculate distally. The remainder of the 
dorsal setae are stout and plumose. The extra-marginal setae are plumose distally. 

The tritosternum is normal with the lacinae strongly pilose. The sternal shield is 
ornamented with a network of ridges and punctures (PL 3, fig. 16). The punctate 
areas are distinct. The first pair of sternal setae (hi) are strongly plumose whereas 
h2 and I13 are simple and blunt apically. The metasternal setae are similar in form 
to setae h2 and I13. The genital shield, strongly punctured, is provided with accessory 
sclerites and a pair of rod-like setae which are spiculate dorsally. The ventri-anal 
shield (approximately 580 x 460/4) is strongly reticulated and punctured. The 
anterior pre-anal setae are simple and the median and posterior setae plumose. The 
para-anals are simple and the post -anal plumose. A pair of well-sclerotized platelets 
lie between the genital and ventri-anal shields. The met apodal shields are well 
developed. The peritreme and peritrematal shield are normal. 

The four pairs of ventral setae on the gnathosoma are simple with the external 
posterior rostrals about one-eighth the length of the internals. The former lie well in 
advance of the latter. The ventral groove is provided with five rows of denticles. 
The tectum consists of a single process, bifurcate distally (Text-fig. 57) . The chelicerae 
are strongly chelate-dentate with the fixed digit bearing two well-developed teeth 
and a strong pilus dentilis (Text-fig. 58). The movable digit is also bidentate. 

Leg I (approx. 480/1) with the tibia (230-240/4) considerably shorter than the 
tarsus (297-320/^). 

Male. The chaetotaxy and ornamentation of the dorsal shield is essentially the 
same as in the female except for a characteristic densely punctured area surrounding 
the base of setae L6. In the majority of the males examined the dorsal shield was 
strongly attentuated in the posterior third. 

The sterniti-genital shield is poorly ornamented with a network of ridges and 
punctures and its posterior margin is slightly concave. The five pairs of setae on the 
shield are all plumose distally. The ventri-anal shield (approximately 560 x 350/1) 
is provided with a polygonal network of ridges and numerous punctations. The 
pre-anal setae and the post-anal seta are plumose in their distal half. The para-anals 
are simple. The interscutal membrane between the sterniti-genital and ventri-anal 
shield is without sclerotized platelets. The metapodal shields are well developed. 

The structure of the venter of the gnathosoma is basically the same as in the female. 
The tectum may be bifurcate or more extensively divided distally (Text-fig. 59). The 
fixed digit of the chelicera is tridentate and the movable bidentate. The spermato- 
phoral process is about one-third the length of the movable digit (Text-fig. 60). 

Femur, genu and tibia of leg II (Text-fig. 61) and the trochanter and femur of leg 
IV (Text-fig. 62) are spurred. 

Dimensions. Male : 1,420-1,520/4 in length, 850-900/4 in breadth. Female : 
1450-1650/4 in length, 790-900/4 in breadth. 

Habitat and distribution. Common in wrack and tidal debris above high- 
water mark on the seashore. Also recorded from salt marshes (Falconer, 1923). 
Widely distributed in Britain, 



BRITISH MITES — SUBFAMILY MACROCHELINAE TRAGARDH 
SPECIES INCERTAE SEDIS 

Macrocheles (Nothrholaspis) pannosus Hull 

Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15 : 211, fig. 

It is possible that this species is a small form of M. plumiventris. 

Dimensions. Female : 900/^ in length. 

Habitat and locality. " In manure, West Allendale." 

Macrocheles (Nothrholaspis) nemoralis Hull 

Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15 : 213, fig. 

This species is possibly a synonym of M. penicilliger. 
Dimensions. Female : S10/1 in length, 500/^ in breadth. 
Habitat and Locality. " West Allendale ; habitat unknown." 

Macrocheles (Nothrholaspis) parmulatus Hull 

Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15 : 214, fig. 

Dimensions. Female : 1,200/^ in length. 

Habitat and locality. " West Allendale ; habitat unknown." 

Macrocheles (Monoplites) palustris Hull 

Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15 : 216, fig. 

Dimensions. Male : 1,100/^ ; female, 1,200/^. 

Habitat and locality. " In sphagnum on the moors, West Allendale." 

Macrocheles (Monoplites) tardior Hull 

Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15 : 216, fig. 

Dimensions. Female : 1,000/t in length. 

Habitat and Locality. " Oxfordshire (R. S. Bagnall). Habitat not stated." 



Genus GEHOLASPIS Berl. s. lat. 

Geholaspis Berlese, A. (1918). Redia, 13 : 145. 

This genus has been revised by Valle (1953) and may be defined as follows : 

Dorsal shield with twenty-eight pairs of setae and twenty-two pairs of " pores ". 

Sternal shield with three pairs of setae ; metasternal shields free or fused with the 

endopodal shields. Ventri-anal shield in the female with five pairs of pre-anal setae. 

Chelicerae normal length with a few teeth or conspicuously elongate and multi- 



42 BRITISH MITES — SUBFAMILY MACROCHELINAE TRAGARDH 

dentate Tectum with median process only. Other characters as in Macrocheles 
Latr. 

Type : Gamasus longispinosus Kramer, 1876. 

Valle (loc. cit.) subdivided the genus into three subgenera of which the following 
two are represented in the British fauna : 

1. Seta Ml long and simple, and extending beyond the bases of D2 by more than half 
their length (fig. 63). Ratio of the length of the dorsal shield to the length of the 
movable digit of the chelicera varies between 6*8 and 10-4 ; movable digit with less 
than 5 teeth ........... Geholaspis s. str. 

Seta Ml short, simple or plumose, and not or scarcely extending beyond the bases of 
seta D2 (fig. 66). Ratio of the length of the dorsal shield to the length of the 
movable digit of the chelicera varies between 3*7 and 4*5 ; movable digit with more 
than five teeth Longicheles Valle. 

The subgenus Geholaspis is represented in Britain by the type species only, and 
Longicheles by two species, namely, G. (L.) mandibularis (Berl.) and G. (L.) longulus 
(Berl.). 

Subgenus GEHOLASPIS Berl. s. str. 

Geholaspis (Geholaspis) longispinosus (Kramer). 

Gamasus longispinosus Kramer, P. (1876). Arch. Naturgesch. 42 : 100, fig. 

Holostaspis longispinosus, Berlese, A. (1887). Acari, Myriapoda et Scorpiones, etc., fasc. 44, 

No. 1, figs. 
? Macrocheles castaneus Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15 : 212. 

Female. The dorsal shield is minutely punctured all over and reticulated in its 
posterior half . It carries twenty-eight pairs of setae and twenty- two pairs of " pores " 
distributed as in Text-fig. 63. Setae Di are strongly plumose ; D2, D5-D8, Mi, M5 
and M6, and Li are simple. The remainder of the dorsal setae are pilose. The extra- 
marginal setae are similar in structure to the marginal series. 

The tritosternum is normal with the lacinae pilose. The sternal shield is ornamented 
with a network of ridges and punctures (PI. 4, fig. 17). The three pairs of sternal 
setae and the metasternals are simple. The genital shield is broad and strongly 
sculptured. The genital setae are simple and the accessory sclerites well-developed. 
The large ventri-anal shield (440-440//; in length and 465-5 io/^ in breadth) is broader 
than long and bears five pairs of pre-anal setae in addition to the three setae normally 
associated with the anus. The relative lengths of these setae are shown in Text-fig. 4. 
The surface of the shield is evenly reticulated. The met apodal shields are small and 
weakly sclerotized. The peritrematal shield is free in its posterior half. 

Ventrally, the gnathosoma bears the normal four pairs of setae. The external 
posterior rostrals lie well in advance of the internals. The ventral groove is provided 
with six transverse rows of denticles. The corniculi are long and slender. The tectum 
comprises a median process arising from a denticulate base (Text-fig. 64). The process 
is provided with shoulder-like projections of varying size and shape about a third of 
the distance from its base ; distally it is divided into two or three pointed branches. 



BRITISH MITES — SUBFAMILY M ACROC HELIN AE TRAGARDH 



43 



The chelicerae are strongly developed and dentate ; the dentition of the digits is 
shown in Text-fig. 65. The ventral setae, omitted from the figure, are well developed. 

Tibia I (approx. 84^) is considerably shorter than tarsus I (approx. 133/0- 

Male. This sex does not appear to have been described. 

Dimensions. Length, 870-970^ ; breadth 590-710/t. 






Figs. 63-65. Geholaspis (Geholaspis) longispinosus (Kramer), female. Fig. 63, dorsal 
shield. Fig. 64, tectum. Fig. 65, chelicera. 



Habitat and locality. This species is widely distributed in Europe (Valle, 1953) 
and relatively common in moss, litter and humus. It has been recorded from North 
Wales by Hull (1918) and from Ireland by Halbert (1915). The writers have examined 
specimens from a number of localities in England, Wales and Scotland. 

The figures of the dorsal and ventri-anal shields of Geholaspis longispinosus 
(Kramer) given by Valle (1953, figs. I and V (2)) do not appear to refer to this species 



44 BRITISH MITES — SUBFAMILY MACROCHELIN AE TRAGARDH 

but to Geholaspis forolivensis Lombardini, 1943, which he considers to be a subspecies 
of longispinosus. In forolivensis the lateral and marginal series of dorsal setae are 
strongly plumose as in mandibularis, and the shape of the ventri-anal shield and the 
relative lengths of the pre-anal setae do not conform with those in longispinosus. 



Subgenus LONGICHELES Valle 
The two British species of this subgenus may be separated as follows : 

1. Setae Mi simple ; a simple seta situated between plumose setae L3 and L4 ; dorsal 

shield less than 650/z in length ...... G.(L.) longulus (Berl.) 

-. Setae Mi strongly plumose ; without simple seta between plumose setae L3 and L4 

(Text-fig. 66) ; dorsal shield about 750^ in length . . G.(L.) mandibularis (Berl.) 



Geholaspis (Longicheles) longulus (Berl.) 

Holostaspis longulus Berlese, A. (1887). Acari, Myriapoda et Scorpiones, etc., fasc. 43, no. 9, 

figs. 
Geholaspis {Longicheles) longulus, Valle, A. (1953). Redia 38 : 351, figs. 

The key characters given above are based on the description and figures of this 
species by Valle (1953). The writers have not examined this species, which is recorded 
by Halbert (1915) from a number of localities in Ireland and by Hull (1918) from 
Ninebanks, Northumberland. 



Geholaspis (Longicheles) mandibularis (Berl.) 

Holostaspis mandibularis Berlese, A. (1904). Redia, 1 : 263. 

? Macrocheles minimus Hull, J. E. (1918). Trans. Nat. Hist. Soc. Northumb. n.s. 5 : 73, fig- 

Geholaspis (Longicheles) mandibularis, Valle, A. (1953). Redia 38 : 344, figs. 

Female The dorsal shield is densely covered with minute tubercles and bears 
twenty-eight pairs of setae, of which twenty- two pairs are strongly plumose. The 
distribution of setae in the posterior half of the shield shows some variation in the 
material examined. The normal chaetotactic pattern is shown in Text-fig. 66 and 
a variant in Text-fig. 67. The lateral margins of the shield are conspicuously ser- 
rated. The vertical setae lie in close proximity to each other ; setae Mi are strongly 
plumose. The extra-marginal setae are plumose and increase in length towards the 
posterior end of the idiosoma. 

The lacinae of the tritosternum are long and pilose. The sternal shield is ornamented 
with strong ridges and tubercles ; the three pairs of sternal setae are simple. The 
elongate metasternals each carry a simple seta. The genital shield is ornamented 
with punctate lines ; the genital setae are simple. The ventri-anal shield is evenly 
reticulated and tuberculated, and is provided with five pairs of pre-anal setae of 
which the external posteriors are pilose. The shape of the shield and the length of 
the setae show considerable variation in the specimens examined (cf. Text-figs. 



BRITISH MITES — SUBFAMILY MACROCHELINAE TRAGARDH 45 





Figs. 66-67. Variation in the chaetotaxy of the dorsal shield of Geholaspis (Longicheles) 

mandibularis (BerL), female. 



46 BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 

68-70). The metapodals are elongate and weakly sclerotized. The posterior half of 
the peritrematal shield is free. 

The external posterior rostrals lie well in advance of the internals and the ventral 
groove is provided with five transverse rows of denticles. The corniculi are long and 
slender and extend beyond the middle of the palp femur. The form of the tectum 
is shown in Text-fig. 71. The chelicerae are very long and strongly toothed (Text-fig. 
72). 

Leg I (approx. 6oo/jl in length) with the tibia (approx. ioo/^.) shorter than the 
tarsus (approx. 130/^). 

Dimensions. Female : length, 750-800/^ breadth, 480-500/^ 

Habitat and locality. In litter, moss and compost. Widely distributed in 
Europe (Franz, 1954). 



Genus MACRHOLASPIS Oudemans. 

Macrholaspis Oudemans, A. C. (1931). Ent. Ber. 8, No. 180 : 272. 

This genus was proposed by Oudemans (193 1) for Gamasus opacus C. L. Koch 
and was characterized by the female having only two pairs of setae on the ventri-anal 
shield (" Ventrianaalschild mit 2 paar borstels "). According to Oudemans one of 
these two pairs of setae would be the para-anals and the other the pre-anals. Recently, 
the writers have examined a preparation and an unpublished drawing of opacus from 
the Oudemans Collection at Leiden, and found that the ventri-anal shield of the 
female, the only sex known, has two pairs of pre-anal setae and not one pair as 
Oudemans has stated. Further, Nothrholaspis aciculatus Berl., a common European 
species, was found to be synonymous with Macrholaspis opacus (Koch) Oudemans, 

1931. 

The following is an emended definition of the genus Macrholaspis : 

Dorsal shield strongly attenuated posteriorly and bearing twenty-nine pairs 1 of 
plumose setae. Lateral margins of the shield markedly serrated. Sternal shield with 
three pairs of setae ; metasternals free. Genital shield with accessory sclerites and a 
pair of genital setae. Ventri-anal shield with two pairs of pre-anal setae. Other 
characters as in Macrocheles. 

Type. Gamasus opacus C. L. Koch, 1839. 

Two species are represented in the British fauna, these may be separated as follows : 

1. Dorsal shield with 58 setae ; Mi short and straight (Text-rig. 73). Serrations of the 
lateral margin of the shield small and rounded (Text-fig. 74). Sternal shield with 
numerous punctures. Posterior margin of trochanter IV smooth 

Macrholaspis opacus (C. L. Koch) 

-. Dorsal shield with 59 setae ; Mi long and curved distally (Text-fig. 78). Serrations 
of the lateral margin of the shield considerably longer and tapering (Text-fig. 80). 
Sternal shield with a network of ridges and punctures. Trochanter IV with a strong 
tubercle on its posterior margin ..... Macrholaspis dentatus sp. n. 

1 An additional seta may be present in some species (cf. Macrholaspis dentatus sp. n.). 



BRITISH MITES — SUBFAMILY MACROCHELINAE TRAGARDH 47 




Figs. 68-72. Geholaspis (Longicheles) mandibulavis (Berl.), female. Figs. 68-70, variation 
in the form of the ventri-anal shield. Fig. 71, tectum. Fig. 72, chelicera. 



48 BRITISH MITES — SUBFAMILY MACROCHELINAE TRAGARDH 

Macrholaspis opacus (C. L. Koch) 

Gamasus opacus Koch, C. L. (1839). Deutsch. Crust. Myr. Arach. fasc. 25, t. 24. 

Macrocheles (Nothrholaspis) aciculatus Berlese, A. (1918). Redia 13 : 169 syn. nov. 

Holostaspis terreus, Halbert, J. N. (1915). Proc. Roy. Irish Acad. 31, ii : 66. 

Macrholaspis opacus, Oudemans, A. C. (1931). Ent. Ber. 8, No. 180 : 272. 

Nothrholaspis aciculatus, Willmann, C. (1939). Ark. Zool. 31 A, 10 : 6, figs. 

Macrocheles {Nothrholaspis) terreus, Cooreman, J. Bull. Mus. roy. Hist. nat. Belg. 19, 63 : 21. 

Female. Dorsal shield strongly attenuated posteriorly (Text-fig. 13) and charac- 
teristically ornamented in its anterior third with a polygonal network of minute spines 
{aciculatus !). The lateral margins of the shield are distinctly serrated (Text-fig. 74). 
The vertical setae are strongly developed ; setae Mi are short and plumose (Text-fig. 
75) . The remaining twenty-seven pairs of plumose setae are distributed as in Text- 
fig. 73. The extra-marginal setae are of the same form as the marginal series. 

The tritosternum is normal with the lacinae pilose. The sternal shield is densely 
covered with punctures and bears three pairs of simple setae (PL 4, fig. 18). The 
metasternal setae are also simple. The genital shield is covered with punctures which 
tend to form a network in its anterior two-thirds. Accessory sclerites are well 
developed and the genital setae are plumose. The ventri-anal shield (approximately 
220fi long and 198/1 wide) is oval in outline and, like the dorsal shield, ornamented 
with spinules. The two pairs of pre-anal setae are plumose, the para-anals long and 
simple, and the post-anal seta strongly plumose. The interscutal membrane between 
the genital and ventri-anal is provided with three pairs of platelets, one pair of pore- 
bearing platelets and a pair of plumose setae. The metapodal shields are conspicuous. 
The peritreme and peritrematal shield are normal. 

The gnathosoma bears four pairs of setae ventrally. The ventral groove has five 
transverse rows of denticles. The form of the tectum is shown in Text-fig. 76. Both 
digits of the chelicera are bi-dentate ; the distal tooth on the fixed digit is small and 
difficult to see if the chelicera is not orientated correctly (Text-fig. yy) . The pilus 
dentilis is long and stout. The dorsal seta is comb-like. 

Leg I (530/^) has plumose setae on the femur, genu and tibia. Tibia I (85^) is con- 
siderably shorter than the tarsus (121/*). Legs II to IV measure 550/*, 495/* and 800/1, 
respectively. 

Male. Unknown. 

Dimensions. Female ; length 725-740/* ; breadth 445-470/*. 

Habitat and locality. Widely distributed in Europe. Common is decaying 
wood, moss and litter. 

Macrholaspis dentatus sp. n. 

Female. Dorsal shield attenuated posteriorly as in the preceeding species but 
without spinules. The ornamentation consists of a network of lines and punctures 
(Text-fig. 78). The chaetotaxy of the shield comprises fifty-nine setae arranged as in 
the figure. The verticals are stout and setae Mi long, plumose and curved (Text-fig. 
79). An interesting feature of the chaetotactic pattern is the presence of three 



BRITISH MITES — SUBFAMILY MACROCHELIN AE TRAGARDH 



49 






6K« \ 



75 





Figs. 73-77. Macvholaspis opacus (C. L. Koch), female. Fig. 73, dorsal shield. Fig. 74, 

lateral margin of dorsal shield. Fig. 75, anterior region of dorsal shield. Fig. 76, 
tectum. Fig. jy, chelicera. 

zool. 4, 1. 4 



50 BRITISH MITES— SUBFAMILY M ACROCHELINAE TRAGARDH 

unpaired setae in the dorsal series. The lateral margins of the shield are strongly 
serrated (Text-fig. 80). The extra-marginal setae are similar to the marginals. 

The sternal shield is provided with a network of ridges and punctures and bears 
three pairs of simple setae (PL 4, fig. 19). The metasternal shields are small and free, 
and the setae simple. The genital shield is strongly ornamented with a network of 
ridges and punctures ; the genital setae are plumose. The ventri-anal shield, similarly 







Figs. 78-80. Macrholaspis dentatus sp. nov., female. Fig. 78, dorsal shield. Fig. 79, 
anterior region of dorsal shield. Fig. 80, lateral margin of dorsal shield. 



ornamented, measures iSy/i in length and 145/* in breadth, and bears two pairs of 
plumose pre-anal setae. The para-anals are long and simple, and the post-anal seta 
short and plumose. The interscutal membrane between the genital and ventri-anal 
shields is provided with platelets and plumose setae as in the preceding species. 
The peritreme and peritrematal shield are normal. 

The chaetotaxy and structure of the gnathosoma are essentially the same as in 
opacus. The form of the tectum and the dentition of the chelicerae are also similar. 

Leg I, approximately 525^ in length, has the tibia considerably shorter than the 



BRITISH MITES — SUBFAMILY MACROCHELINAE TRAGARDH 51 

tarsus (tibia, 88/1 ; tarsus, 110/1). Legs II to IV measure 575/4, 560/* and 880/1 in 
length, respectively. 

Dimensions. Length 765/4 ; breadth 430/1. 

Habitat and locality. A single female from humus under bracken in the Leri 
Valley, North Cardiganshire, Wales. Holotype female, 1955.10.22.95. 

This species is characterized by the form of setae Mi, the ornamentation of the 
sternal shield and the shape of the ventri-anal shield. 



Genus HOLOSTASPELLA Berl. 
Holostaspella Berlese A. (1904). Redia 1 : 241 

Dorsal shield with twenty-eight pairs of setae. Vertical setae situated on an 
outgrowth of the dorsal shield. Sternal shield with three pairs of setae ; metasternal 
shields free. Ventri-anal shield with four pairs of pre-anal setae. Gnathosoma normal. 
Leg I in the female armed with stout spurs. Other characters as in Macrocheles Latr. 

Type. Holostaspis (Holostaspella) sculpta Berl., 1904. 

This genus is represented in Britain by one species only. 



Holostaspella ornata (Berl.) 

Macrocheles vagabundus, Oudemans, A. C. (1902). Tijdschr. Ent. 45 : 43, figs. 
Holostaspis ornatus Berlese, A. (1904). Redia 1 : 277. 
Holostaspella ornata Oudemans, A. C. (1931)- Ent. Ber. 8 : 273, syn. nov. 
Holostaspella ornata, Evans, G. O. (1956). Proc. Linn. Soc. Lond. 

Female. Dorsal shield, finely reticulated, bears twenty-eight pairs of setae 
(Text-fig. 81). The verticals are stout and pilose, and are situated on a prominent 
projection (Text-fig. 82). The setae on the shield are minutely pilose as shown in the 
figure. 

The sternal shield is massive and characteristically ornamented with ridges and 
depressions (PL 4, fig. 20). The first pair of sternal setae are strongly plumose ; the 
second and third pair smooth. The metasternals, triangular in outline, each bear a 
simple seta. The large genital shield is richly ornamented ; the genital setae simple. 
Most of the region posterior to coxae IV is occupied by the heavily ornamented 
ventri-anal shield (335 X 315/^), which bears four pairs of pre-anal setae in addition 
to the para-anals and the post-anal seta. The anterior pre-anal is weakly pilose and 
the post-anal plumose. The interscutal membrane is coarsely striated. 

The gnathosoma is normal for the Macrochelinae. The lateral processes of the 
tectum are free (Text-fig. 83). The dentition of the chelicera is shown in Text-fig. 84. 

Leg I with the tibia (121/4) shorter than the tarsus (132/4) . The femur and trochanter 
of leg II are spurred ; the tarsus has a large stout spine ventrally (Text-fig. 85). 

Male. Unknown. 



52 BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 




Figs. 81-85. Holostaspella ornata (Berl.), female. Fig. 81, dorsal shield. Fig. 82, 
anterior region of dorsal shield. Fig. 83, tectum. Fig. 84, chelicera. Fig. 85, leg II 
(ambulacrum omitted). 



BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 53 

Dimensions. Length 924-950/t ; breadth 550-560/^ 

Habitat and locality. This species has previously been recorded from the 
Netherlands (Oudemans, 1902) and Austria (Franz, 1954). The writers have examined 
a single specimen taken from Sphaerocerus sp. captured at Bagley Wood, Berks 
(Coll. O. W. Richards, det. Vitzthum). The above description and figures are based 
on the type from the Oudemans Collection, Leiden. 



SUMMARY 

1. The classification of the Macrochelidae is discussed and keys are given for the 
identification of British species of the Macrochelinae. 

2. The following three new species are described and figured : 

Macrocheles rothamstedensis sp. no v. 
Macrocheles punctoscutatus sp. no v. 
Macrholaspis dentatus sp. no v. 

3. The following species are recorded for the first time from Britain : 

Macrocheles carinatus (C. L. Koch) 
Macrocheles decoloratus (C. L. Koch) 
Macrocheles insignitus (Berl.) 
Macrocheles montanus (Willmann) 
Macrocheles penicilliger (Berl.) 
Macrocheles subbadius (Berl.) 
Macrocheles tardus (C. L. Koch). 
Geholaspis (Longicheles) mandibularis (Berl.) 
Holostaspella ornata (Berl.) 

4. The following species have been relegated to the synonymy : 

Nothrholaspis aciculatus Berl., 1918 syn. of Macrholaspis op acus (C. L. Koch) 

1839. 
Holostaspella ornata Oudemans, 1931, syn. of Holostaspella ornata (Berl.), 1904. 
Macrocheles gladiator Hull, 1918, syn.(?) of Macrocheles plumiventris Hull, 1918. 
Macrocheles plumipes Hull, 1918, syn.(?) of Macrocheles plumiventris Hull, 1918. 
Macrocheles minimus Hull, 1918, syn. of G. {Longicheles) mandibularis (Berl.), 

1904. 
Macrocheles hulli Falconer, 1923, syn. of Macrocheles carinatus (C. L. Koch), 

1839. 
Macrocheles occidentalis Hull, 1925, syn. of Macrocheles submotus Falconer, 

1924. 
Macrocheles gloriosus Hull, 1925, syn. of Macrocheles submotus Falconer, 1924. 
Nothrholaspis fimicola Sellnick, 1932, syn. of Macrocheles plumiventris Hull, 

I925- 
Coprholaspis anglicus Turk, 1946, syn. of Macrocheles glaber (Miiller), i860. 



54 BRITISH MITES — SUBFAMILY M ACROCHELIN AE TRAGARDH 



ACKNOWLEDGMENTS 



We are extremely grateful to Dr. L. van der Hammen for the loan of specimens 
from the Oudemans Collection at Leiden ; to Dr. F. A. Turk for the loan of the type 
of Coprholaspis anglicus Turk ; to Dr. G. Lombardini for comparing some of our 
specimens with the types in the Berlese Collection, Florence and to Mr. M. G. Sawyers, 
British Museum (Nat. Hist.) for the photomicrographs. 



REFERENCES 

Berlese, A. 1882. Gamasidi nuovi e poco noti. Boll. Soc. ent. ital. 14 : 338-352. 

1 882-1 903. Acari, Myriopoda et Scorpiones hucusque in Italia reperta, Portici et Padua. 

Fasc. 1-101. 

1904. Acari nuovi I and II. Redia, 1 : 235-280. 

1918. Centuria quatria di Acari nuovi. Redia, 13 : 1 15-192. 

1 92 1. Indice sinonimico dei generi e delle specie illustrate nei fascicolo 1 a 101. Redia, 

14 : 77-105. 

Cooreman, J. 1943. Note sur la faune des Hautes-Fagnes en Belgique (1) XI. Acariens 
(Parasitiformes) (2). Bull. Mus. roy. Hist. nat. Belg. 19, No. 63 : 1-28. 

Evans, G. O. 1956. An introduction to the British Mesostigmata with keys to familes and 
genera. /. Linn. Soc. Lond. Zool. (in press) 

Falconer, W. 1923. Two British mites new to science and a new subgenus of Macrocheles 
Latr. Naturalist, Lond. : 151-153. 

1924. Macrocheles submotus — a new name for M. cognatus Falcr. (nom. praeocc). 

Naturalist, Lond. : 363. 

Franz, H. 1954. Die Nordost-Alpen. 15. Ordnung Acarina in Spiegel Land-Tierwelt Inns- 
bruck, 1 : 329-452. 

Hermann, J. F. 1804. Memoire apterologique . Strasbourg, pp. vi -f- 144, 9 pis. 

Halbert, J. N. 1915. Clare Island Survey. Acarinida: ii. Terrestrial and marine Acarina. 
Proc. roy. Irish Acad. 3 39 ii : 45-136. 

Hughes, A. M. 1948 .Mites associated with stored food products. London: H. M.S. O., pp. 168. 

Hull, J. E. 1918. Terrestrial Acari of the Tyne Province. Trans, nat. Hist. Soc. Northumb. 
n.s., 5 : 13-88. 

1925. Acari of the family Gamasidae : new and rare British species. Ann. Mag. nat. 

Hist. (9) 15 : 201-219. 

Hyatt, K. H. 1956. A collection of mites from stable manure. Ent. mon. Mag. (m press). 
Koch, C. L. 1836-1841. Deutschlands Crustaceen, Myriopoden und Arachniden, Regensburg. 
Kramer, P. 1876. Zur Naturgeschichte einiger Gattungen aus den Familie der Gamasiden. 

Arch. Naturgesch. 42 : 46-105. 
Muller, J. i860. Insectenepizoen der mahrischen Fauna (Jahr. naturw. sect.) K.K. mdhr. 

schles. Gesell. Ackevbau. Natur.-u. Landes : 157-184. 
Oudemans, A. C. 1901. Bemerkungen iiber Sanremeser Acari. Tijdschr. Ent. 43:129-139. 

1 91 3. Acarologische Aanteekeningen. Ent. Ber. 4, 73 : 2-18. 

1 91 4. Acarologisches aus Maulwurfsnestern. Arch. Naturgesch. 79a, 8 : 108-200. 

1 931. Acarologische Aanteekeningen CIX. Ent. Ber. 8, 180 : 272-280. 

Pereira, C. & Castro, M. P. de. 1945. Contribuicao para o conhecimento da especie tipo 
de Macrocheles Latr. (Acarina) : M. muscaedomesticae (Scopoli, 1772) emend. Arq. Inst. 
Biol. S. Paulo, 16 : 153-186. 

Scopoli, J. A. 1772. Annus V. Historico naturalis. Lipsiae. 



BRITISH MITES— SUBFAMILY MACROCHELIN AE TRAGARDH 55 

Sellnick, M. i 93 i. Zoologische Forschungreise nach den Jonischen Inseln und dem 
Peloponnes von Max Beier, Wien. Acari. S.B. Akad. Wiss. Wien, 140 : 693-776. 

1940. Die Milbenfauna Islands. Goteborg. Vetensk. Samh. Handl. (5) 6b, 14 : 1-129. 

Tragardh, I. 1949. Description of two new genera of Mesostigmata (Acarina) . Aspidilaelaps 

from Samoa and Protoholaspis from Peru. Ent. Medd. 25 : 311—325. 

1952. Acarina collected by the Mangarevan expedition to South Eastern Polynesia in 

1934 by the Bernice P. Bishop Museum, Honolulu, Hawaii. Mesostigmata. Ark. Zool. 
(2) 4 : 45-90. 

Turk, F. A. 1946. Studies on Acari. V : Notes on and descriptions of new and little- 
known British Acari. Ann. Mag. nat. Hist. (11) 12 : 785-820. 

1948. Insecticolous Acari from Trinidad, B.W.I. Proc. Zool. Soc. Lond. 118 : 82-125. 

Valle, A. 1953. Revisione di generi e sottogeneri berlesiana di Acari. Redia, 38 : 316-360. 
Willmann, C. 1939. Die Arthropodenfauna von Madeira . . . etc. XIV. Terrestrische 

Acari (exkl. Ixodidae). Ark . Zool. 31a 10 : 1-42. 

1 95 1. Die hochalpine Milbenfauna der mittleren Hohen Tauern insbesondere 

des Grossglockner Gebietes. (Acari). Bonner Zool. Beitr. 2 : 141-176. 




6 APR 19SC 



EXPLANATION OF PLATES 
PLATE i 
Sternal, genital and ventri-anal shields of the females of : 



Fig. i 
Fig. 2 
Fig. 3 
Fig. 4 
Fig. 5 
Fig. 6 



Macrocheles muscaedomesticae (Scopoli). x 95. 
Macvocheles rothamstedensis sp. nov. x 153. 
Macrocheles glaber (Miiller). x 115. 
Macrocheles punctoscutatus sp. nov. x no. 
Macrocheles subbadius (Berl.). X 160. 
Macrocheles insignitus Berl. x 218. 



Bull B.M. {N.H.) Zool 4, 1. 



PLATE I 




b 



PLATE 2 
Sternal, genital and ventri-anal shields of the females of : 

Macrocheles merdarius (Berl.). x 199. 
Macrocheles montanus (Willmann). x 100. 
Macrocheles carinatus (C. L. Koch). x 100. 
Macrocheles penicilliger (Berl.). x no. 
Macrocheles submotus Falconer. x 90. 
Macrocheles tardus (C. L. Koch). x no. 



JhlG. 


7- 


Fig. 


8. 


Fig. 


9- 


Fig. 


10. 


Fig. 


n. 


Fig. 


12. 



Bull. B.M. (N.H.) Zool. 4. 1 



PLATE II 




PLATE 3 
Sternal, genital and ventri-anal shields of the females of 

Fig. 13. Macrocheles decolor atus (C. L. Koch). x 

Fig. 14. Macrocheles matrius Hull. X 112. 

Fig. 15. Macrocheles plumiventris Hull. x 74. 

Fig. 16. Macrocheles superbus Hull. x 70. 



Bull. B.M. (N.H.) Zool. 4, 



PLATE 111 







16 



PLATE 

Sternal, 

Fig. 17. Geholaspis {Geholaspis) longispinosus (Kramer). x ioo. 

Fig. 18. Macrholaspis opacits (C. L. Koch). X 153. 

Fig. 19. Macrholaspis dentaius sp. nov. x 131. 

Fig. 20. Holostaspella ornata (Berl.). x 123. 



Bull. B.M. (N.H.) Zoo!. 4, 1. 



PLATE IV 




i i AUGT956 THE EVOLUTION OF 

RATITES 



SIR GAVIN de BEER 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 4 No. 2 

LONDON: 1956 



THE EVOLUTION OF RATITES 




BY 



SIR GAVIN de BEER, F.R.S. 



Pp. 57-70 ; Plates 5-9 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 4 No. 2 

LONDON: 1956 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 4, No. 2 of the Zoological series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued August, 1956 Price Ten Shillings 



THE EVOLUTION OF RATITES 

By Sir GAVIN de BEER, F.R.S. 

Director, British Museum (Natural History). 

SYNOPSIS 

Adequate knowledge of the structure of Archaeopteryx now enables a comparison to be made 
between it and the Carinates. In the latter the structure of the wing, the tail, and the cerebellum 
can be shown to be adaptations to flight. Since these same adaptations are found in Ratites, 
they would be inexplicable unless the Ratites were descended from flying birds. The palate 
of the Ratites is not primitive but neotenous, and represents an early stage through which the 
palate of many Carinates passes during development. Other neotenous features of the Ratites 
are the plumage and the persistence of sutures between the bones of the skull. 



INTRODUCTION 

In the first edition of the Origin of Species (1859, P- 1 34) Darwin wrote : "As 
Professor Owen has remarked, there is no greater anomaly of nature than a bird that 
cannot fly ; yet there are several in this state." There was a touch of irony in 
making this quotation, for as is well known, Owen's views on evolution were uncertain 
and equivocal, and the very existence of flightless birds was inescapable evidence of 
descent with modification from the " archetype " of birds. Darwin himself 
continued : " We may believe that the progenitor of the ostrich genus had habits 
like those of the bustard, and that, as the size and weight of its body were increased 
during successive generations, its legs were used more, and its wings less, until they 
became incapable of flight." 

A few years later, in his treatise on The Anatomy of Vertebrates (1866, 2, 12.) 
Owen put forward the view that the Cursores or Ratites were not " a natural order ; 
some of its exponents have demonstrably closer affinities to other groups of which 
they are wingless members." Further on (loc. cit. : 43), Owen referred to the Ratites 
as " those birds in which the power of flight is abrogated ". For a man who did 
not believe unreservedly in evolution, this was about as near as he could get to the 
view that the Ratites are descended from flying birds, and he even supplied the 
explanation of such a descent (loc. cit. : 12) : "by the arrested development of the 
wings unfitting them for flight ". 

The view of the degenerate nature of the Ratites has been supported by M. 
Fiirbringer (1888), T. J. Parker (1892), W. P. Pycraft (1901), R. Broom (1906), J. E. 
Duerden (1920), E. Stresemann (1927-34), W. K. Gregory (1935), and many others ; 
and it might have been thought that the evolution of flightless birds from flying 
birds was generally accepted. Nevertheless the hypothesis has been put forward 
that the structure of the Ratites is in many respects so primitive that they must have 

ZOOL. 4, NO. 2. 5 



60 THE EVOLUTION OF RATITES 

branched off from the main stem of bird evolution before the power of flight was 
acquired. B. Lindsay (1885), R. S. Wray (1887), A. C. Chandler (1916) and J. C. 
Ewart (1921) are of this opinion, but the foremost exponent of this view has been 
P. R. Lowe (1928, 1935), with whom M. Friant (1945, 1946) has expressed agreement. 

It has been known for a long time that the Ratites show a number of characters 
which have been considered as primitive. Among these is the palate, on which T. H. 
Huxley (1867) based a system of classification of the birds in which the dromaeogna- 
thous type, characteristic of the Ratites, was regarded as the most primitive. 
Pycraft (1900, 1901) who extended Huxley's observations, summed up (1901 : 343) 
the situation as he saw it in the words : " The contention that the Struthious 
(Palaeognathine) palate is of a more ancient type than the Neognathine is admitted 
by all." The term palaeognathine or palaeognathous is equivalent to dromaeo- 
gnathous, while neognathine or neognathous includes all the other categories of 
Huxley's classification. 

This argument has been adopted by Lowe, in whose view the Ratites show " the 
primitive palate from which the neognathous palate characteristic of the modern 
or flying birds was obviously derived ". To this Lowe has added six further argu- 
ments, all in support of his view that the ancestors of Ratites never flew. They are : 

1. The " primitive " disposition of the muscles. 

2. The fact that " all the feathers borne by the adult ostrich or by any other 
struthious form whether they are situated on the wing, or body generally, are 
nothing more than down, or modified down," and being juvenile structures are 
he thinks therefore ancestral and primitive. 

3. The absence of the rudiment of the clavicle in the embryo of the ostrich, 
which, in his view, is proof that the ostrich decended from ancestors which had 
lost the clavicle, and therefore not from flying birds in which the clavicle is 
preserved. 

4. The persistence of the sutures in the skull of the ostrich, in which it 
resembles the condition of the ancestral reptiles and differs from that of flying 
birds in which the bones of the skull are firmly fused in the adult. 

5. The obtuse angle subtended between the coracoid and the scapula, which 
resembles the condition of the ancestral reptiles and differs from that of flying 
birds where this angle is more or less acute. 

6. The similarity between the bones of the hand of the ostrich and those of 
a dinosaur such as Ornitholestes. 

With perfect logic, Lowe contended that if, as he believed, the Ratites were 
descended from birds which had never acquired the power of flight, then it must 
follow that Archaeopteryx could not have been in the line of ancestry of birds, but 
must have been an independent offshoot from the reptiles. 

The detailed knowledge now available of the structure of Archaeopteryx (de Beer, 
19546) can be used to test this hypothesis. Two related problems are involved : 
the evolution from reptiles of birds in general, and the evolution of Ratites in 
particular. These problems can be solved by finding the answers to the following 
three questions : 



THE EVOLUTION OF RATITES 61 

i. Is Archaeopteryx on the line of evolution from reptiles to birds? 

2. Are there any characters by which modern flying birds differ from Archaeopteryx 
which can be attributed with certainty to adaption to active flight? 

3. Are these characters also shown by the Ratites? 



ARCHAEOPTERYX AND THE ANCESTRY OF BIRDS 

If Archaeopteryx was the product of an independent line of evolution from the 
reptiles, unrelated to the stock which gave rise to birds, it then becomes necessary to 
believe that the feathers of Archaeopteryx and the feathers of all other birds were 
independently evolved. The identical details of structure which the feathers show 
involve the quill, the vane formed of barbs, held together and parallel with one 
another and yet capable of being torn apart, the proportions between the vane and 
the quill and between the proximal and distal portions of the vane. In all these 
respects, the structure of the feather in Archaeopteryx and in modern flying birds is 
so exactly identical that it is impossible to believe that they were independently 
evolved. 

But this is not all. In addition to the feathers themselves, there is the manner in 
which they are arranged on the wing, the differentiation between larger feathers or 
remiges and smaller feathers or coverts, and the further differentiation of the remiges 
into primaries, borne on the wrist-joint and hand, and secondaries borne on the 
forearm. Here again the conditions are identical in Archaeopteryx and in modern 
flying birds. It follows that the view that Archaeopteryx is not related to the modern 
birds is completely untenable. 

Granting that Archaeopteryx represents an example of an early stage in the evolu- 
tion of feathered organisms away from the reptiles, it may still be asked whether 
Archaeopteryx is ancestral to modern birds. The remarkable mosaic of reptilian 
and avian characters that Archaeopteryx shows has been discussed elsewhere (de Beer, 
1954a). The conclusion to be drawn is that Archaeopteryx is a rara avis among 
fossils in that it is possible to say that nothing is known, either by way of structures 
which it possesses or does not possess, or of the time-relations of its occurrence, which 
might disqualify it from being regarded as a true ancestor of modern birds. As G. G. 
Simpson (1936 : 92) has said, " every difference between Archaeopteryx ... on 
one side and true reptiles of possible ancestral type, especially the Pseudosuchia, 
on the other, is definitely in the direction of true birds ". 

If, as H. Steiner (1918, 1956) believes, Archaeopteryx was aquintocubital, it 
would provide yet another proof that it was ancestral to modern birds. 



CARINATES AND ADAPTATIONS TO FLIGHT 

Accepting the fact that Archaeopteryx is a mile-stone on the road from reptiles to 
modern birds and represents the type of structure from which Carinates evolved, 
attention may be turned to the question whether any of the differences observable 
between Archaeopteryx and the modern flying birds or Carinates can with certainty 
be ascribed to adaptation to flight. That the flying bird is highly adapted to its 



62 THE EVOLUTION OF RATITES 

mode of life is a commonplace of biological expression, and in the case of some 
structures it is easy to prove it. Attention will here be confined to the carpo- 
metacarpus, the pygostyle, and the cerebellum. The keel on the sternum is 
deliberately omitted from the discussion since its absence in the Ratites is the basis 
of their diagnosis, and the question at issue is whether this absence is primitive or 
specialized. 

The carpometacarpus is the product of fusion between the distal carpals and the 
three metacarpals, the 2nd and 3rd of which are fused again at their distal extremities. 
The result is a structure providing a light and resilient yet firm basis for the attach- 
ment of the primary remiges. It is absolutely characteristic of modern birds and 
found nowhere else. 

In Archaeopteryx the forelimb skeleton consists of proximal carpals, the radiale and 
ulnare which remain more or less separate, distal carpals fused together and to the 
base of the third metacarpal, and separate and independent 1st and 2nd metacarpals. 
Archaeopteryx was unable to do much more than glide, and as Dr. H. W. Parker has 
remarked to me, the air-pressure on the feathers of its wings must have been lower 
than in an actively flying bird with the same ratio of wing-area to mass, because 
Archaeopteryx was unable to maintain itself in the air continuously against the pull 
of gravity. The carpometacarpus of the Carinate is without doubt an adaptation 
to flight by enabling the wing to exert and withstand greater pressure. 

There is one further feature of the wing of Carinates that calls for notice, and that 
is the presence of a small number of feathers attached to the first digit of the hand, 
forming a " bastard wing ". These few feathers add nothing to the weight-bearing 
power of the wing, yet they perform a function of capital importance in flight for 
they serve like the slotted wing of a modern aircraft to maintain a slip-stream of air 
and prevent stalling. The " bastard wing " is a beautiful adaptation to flight. 

In Archaeopteryx the tail is very long, as long as the rest of the body, and its 
skeleton consists of 20 elongated separate vertebrae, to each of the hinder 15 of which 
correspond a pair of rectrices, quill-feathers, which form an oblong and elongated 
air-resisting surface. In Carinates the tail is very short, consisting of about a dozen 
flattened vertebrae, the hindmost half-dozen or so of which are fused together, 
giving rise to a pygostyle. The rectrices, to the number of a dozen pairs, are disposed 
transversely. A masterly analysis and comparison of the conditions in the tail of 
Archaeopteryx and Carinates has been given by H. Steiner (1938). 

It has been pointed out by J. Maynard Smith (1952) that the structure of the most 
primitive flying animals is one that imparts aerodynamic stability. That is to say 
that they are of a shape such that when in " flight " through the air, they are able to 
maintain the direction of their progress without muscular intervention and 
compensatory movements. In other words, such animals are gliders, and the 
structure of the skeleton, wings, and tail of Archaeopteryx is just such as would have 
enabled it to glide with stability, but not fly actively. The perfection of the power 
to fly has involved the development of the ability to perform mechanically unstable 
flight-movements, such as rapid pitching, yawing, and banking, for which a reduction 
of the long axis of the animal is essential. The pygostyle of the Carinates is without 
doubt an adaptation to flight. 



THE EVOLUTION OF RATITES 63 

The cerebellum of Archaeopteryx is best characterized negatively by saying that 
it is small and does not overlap forwards over the midbrain. In other words, the 
brain of Archaeopteryx is similar to that of reptiles. In Carinates, on the other hand, 
the cerebellum is so large that it expands forwards as a median and unpaired structure 
over the dorsal surface of the midbrain, which it presses downwards, and so the 
cerebellum comes into contact with the hinder part of the cerebral hemisphere. The 
result is that the cerebellum of Carinates hides the optic lobes, whereas the latter 
structures are plainly visible in Archaeopteryx. 

The cerebellum has been defined by Sherrington as the head-ganglion of the 
proprioceptive system. As L. Edinger (1912 : 300) has shown in Columba, among 
the most important sources of impulses conducted to the cerebellum are the organs 
of balance in the semi-circular canals of the ear, which respond not only to changes 
in static conditions, but also to changes in the dynamic conditions of the organism 
caused by alterations in speed and direction of motion. In the case of Carinates, 
as flying birds the possibilities of direction of motion are greatly increased by the 
introduction of the vertical dimension. At the same time, the performance of 
flight requires high speed of adjustment and compensatory movements, not only in 
actual flight but in landing on small objects. As J. Z. Young (1950 : 455) has said, 
there is reason to think that the large size of the cerebellum in flying birds is 
connected with the precise control of movement in all planes of space during flight. 

The view that the cerebellum of Carinates is an adaptation to flight is confirmed 
by the conditions in the pterosaurs. There, as T. Edinger (1941 : 678) has shown, 
there is " a cerebellum thrust forward above the midbrain to adjoin the forebrain as 
in birds ; obviously this is one of the characters distinguishing all pterosauria from 
the other reptiles ". There can be no doubt that the parallel development and large 
size of the cerebellum in pterosaurs and in birds, by which they both differ from all 
non-pterosaurian reptiles, are due to the same cause : adaptation to flight. 

Having now established that the carpometacarpus, the " bastard wing ", the 
pygostyle, and the large size of the cerebellum of Carinates are adaptations to flight, 
attention may be turned to the conditions in Ratites in respect of these structures. 

ADAPTATIONS TO FLIGHT IN RATITES 

The skeleton of the wing of Ratites is built on identically the same plan as that of 
the Carinates. B. W. Tucker (1938a : 224) has stressed the similarity not only in 
the points of fusion between the various elements which go to make up the carpometa- 
carpus in both Carinates and Ratites, but also subtle points, such as the curvatures 
of the 2nd and 3rd metacarpals and the expansion of the basal phalanx of the 3rd 
digit. 

H. Steiner (1949 : 367) has studied the wings of Carinates and Ratites by means of 
X-rays and concludes that it " lasst sich ohne jeden Zweifel feststellen, dass genau 
die gleichen Eigentumlichkeiten zu beachten sind. Ausgehend von irgend einem 
Carinatenflugel kann ausserdem iiber den Fliigel von Rhea und Struthio bis zu jenem 
von Casuarius eine zunehmende Verkummerung verfolgt werden, welche iiber die 
Zustande, wie sie bei Dromaeus und Apteryx angetroffen werden, bis zur vollstandigen 



64 THE EVOLUTION OF RATITES 

Reduktion des Fliigels bei den ausgestorbenen Riesenstraussen Aepyornis und 
Dinornis gefuhrt hat ". 

There can be no doubt that the skeleton of the wing in the Ratites shows features 
associated with adaptation to flight which are explicable only on the view that they 
were inherited from ancestors which flew. 

The skeleton of the tail in Ratites has been studied by W. Marshall (1873) and 
referred to by W. K. Gregory (1935), but otherwise has not attracted much attention. 
It is composed of a varying small number of vertebrae which in some forms decrease 
in size caudally and taper out. But in the ostrich there is a structure composed of 
the fusion of the terminal vertebrae which undoubtedly constitutes a pygostyle. It 
is shown in PI. 5 in comparison with a Carinate pygostyle. Since this structure in 
Carinates is certainly associated with the power of flight, its presence in a Ratite is 
inexplicable unless the ancestors of Ratites also flew. This has also been pointed out 
by Gregory. 

The cerebellum of birds has been subject to an exhaustive study by S. Ingvar 
(1918), the results of which show that the large size of the cerebellum in Carinates is 
matched by a similar large size in Ratites. Not only does the cerebellum of the 
ostrich, for instance, project forwards over the dorsal surface of the mid-brain towards 
the cerebral hemisphere, but it shows the same arbor vitae structure as the cerebellum 
of a Carinate when seen in sagittal section. In Plate 6 are shown the brain of 
Archaeopteryx in side view, and sagittal sections through the brains of Rhea and Tringa. 
It is clear that the structure of the cerebellum is the same in the Ratites as in the 
Carinates ; and if its structure in the latter is an adaptation to flight, its structure 
in the former is inexplicable unless the Ratites were descended from flying birds. 

On all three counts, the evidence is conclusive that the Ratites must have evolved 
from flying birds. It remains now to consider a few further points which receive 
ready explanation on this view, and to refute the grounds on which Lowe thought 
that the Ratites were primitively flightless. 

It has been shown above that the skeleton of the wing of the Ratites bears evidence 
of adaptation to flight. It may be added that in one form, the Rhea, there is still to 
be seen a trace of the differentiation between primary and secondary remiges, as 
shown in Plate 7. This distinction, which already exists in Archaeopteryx would be 
meaningless unless the Rhea's ancestors had been capable of flight. Even more 
remarkable is the presence in the Rhea of feathers on the 1st digit forming a " bastard 
wing ", an adaptation evolved in Carinates which results in the maintenance of the 
slip-stream in flight. 

The curious phenomenon of diastataxy or aquintocubitalism, the absence of the 
5th secondary remex from the row of flight feathers in the wing, has long been a 
puzzle. Its most probable explanation has been provided by H. Steiner (191 8) who 
has shown in a brilliant and exhaustive series of studies that it is associated with the 
peculiar method of folding the wing, the ulnar flexure, adopted by birds. When a 
bird folds its wings, the hand is moved sideways relatively to the forearm through 
an angle of almost 180 . The development of this new type of movement affected 
the feather-rudiments in the skin at the point of flexure and dislocated them in such 
a way that the rudiment which would have given rise to the 5th secondary remex is 



THE EVOLUTION OF RATITES 65 

displaced, and, instead, develops into the 5th major covert, leaving a gap in the 
series of secondaries. Steiner has shown conclusively that in the Carinates the 
aquintocubital condition is primitive, and that the presence of the 5th secondary 
remex, which is found sporadically in some members of nearly all groups of birds, is 
due to a secondary readjustment. Be that as it may, it is clear that the phenomenon 
of aquintocubitalism is intimately associated with the structure and arrangement of 
the remiges in a flying wing. It is therefore remarkable that a vestige of the aquinto- 
cubital condition is found in the wing of the young Apteryx (Steiner 1918 : 434) which 
thereby is shown to possess a structure characteristic of primitive Carinates and 
which could not have been independently evolved. Professor Steiner has kindly 
informed me that he has evidence that other Ratites also are aquintocubital. 

Further, there is another line of evidence relating to the loss of the power of 
flight of Apteryx. R. Broom (1947 : 49) has ingeniously shown that as New Zealand 
has had no land connexions with any other continent since early Jurassic times, and 
as the centre of evolution of birds exemplified by Archaeopteryx was situated in the 
Palaearctic continent in middle Jurassic times, the ancestors of Apteryx could not 
have reached New Zealand unless they flew thither. 

THE NEOTENY OF RATITES 

Reverting now to the reasons on which Lowe sought to base the view that the Ratites 
were primitive birds whose ancestors had never flown, one : the similarity between 
the hand of the ostrich and that of the dinosaur, has been dismissed as invalid. 
Tucker (19386) has shown that such resemblances as there are between them are 
only superficial and without significance. Another : the angle between the coracoid 
and the scapula, can be shown to be due to the reduction of the length of the pectoral 
muscles in the Ratites ; for it is the lengthening of the coracoid in the Carinates 
which is responsible for the acuteness of the angle between the coracoid and the 
scapula; and the length of the coracoid may be regarded as an adaptation to flight 
since it is associated with the lengthening of the pectoral muscles. 

Whether the disposition of the muscles in the Ratites is " primitive ", as Lowe 
has contended, is a matter for argument ; but what is no matter for argument is the 
explanation of the presence in the Ratites of nestling-down, permanent sutures 
between the bones of the skull, and the dromaeognathous structure of the palate. 
All these are demonstrably the result of neoteny or the secondary retention of 
features which were juvenile in the ancestors of the Ratites. 

To begin with the feathers. It is well known that the down-feathers, nestling- 
down or neossoptyles, are nothing but the fluffed-out distal ends of the rudiments of 
the adult feathers or teleoptyles. In Carinates, particularly those in which the young 
are nidifugous and have a " chick " stage, the nestling-down is well developed, and 
it owes its flufnness to the fact that the barbs have no hooks and therefore the 
feathers form no vanes. This nestling-down is subsequently discarded when the 
adult feathers or teleoptyles take the place of their former distal extremities the 
neossoptyles. That the Ratites are neotenous in retaining their " ostrich feathers " 
or nestling-down throughout life is admitted by Lowe himself (1935 : 420) : " So far 



66 THE EVOLUTION OF RATITES 

as their feather covering is concerned the Struthiones are big, overgrown chicks. 
They are the " Peter Pans " of the avian world. They have never grown up." 

The same phenomenon of neoteny is responsible for the retention of the sutures 
between the bones of the skull in the ostrich. In the Carinates, the sutures between 
the bones are present in the young stages, but they are obliterated in the adult skull, 
which is a structure of great solidity, in all probability adapted to the necessity for 
withstanding the mechanical stresses consequent upon active flight. In retaining 
the sutures between the bones of the skull the ostrich, having lost the power of flight, 
shows a secondary return to the juvenile condition of the ancestral flying bird, and, 
of course, the ancestral reptile. 

The inclusion of the dromaeognathous or palaeognathous palate among the 
neotenous features of the Ratites, with the implication that it is the result of a 
secondary retention of an ancestral juvenile condition, may appear surprising in 
view of the selection of this very structure by T. H. Huxley as the basis for his 
classification of birds, and his view that the dromaeognathous type was primitive. 
Nevertheless, the evidence is quite clear, as W. P. Pycraft (1900, 1901) has shown, 
although he did not realize its significance. The so-called palaeognathous palate is 
an arrested stage in the development of the neognathous palate. Precisely the same 
conclusion was reached by S. McDowell (1948) on other grounds, namely the impos- 
sibility of giving a definition of the palaeognathous palate applicable to all Ratites 
and tinamus and excluding all Carinates (except tinamus) because of its great 
variation. 

The essential feature of Huxley's dromaeognathous and Pycraft's paleognathous 
palate is the fact that the pterygoids extend forwards and make contact with the 
hinder ends of the pre vomers, while the palatines lie further to the side. In Huxley's 
schizognathous and aegithognathous types, or Pycraft's neognathous palates, the 
usual condition in the adult is that the pterygoids do not make contact with the 
prevomers, but are separated from them by the palatines with which the pterygoids 
make a joint. But Pycraft's remarkable discovery, to which insufficient attention 
has been paid, was that in the development of many " neognathous " birds the 
palate passes through a " palaeognathous " stage in which the pterygoids actually or 
nearly come into contact with the prevomers ; but the anterior ends of the pterygoids 
then become detached from the remainder of these bones, and, instead, become 
attached to the hinder ends of the palatines, where they give rise to the so-called 
" mesopterygoid " elements of W. K. Parker (1875, 1876, 1877, 1879), an d the 
" hemipterygoid " of Pycraft. Between the detached anterior portion and the 
remainder of the pterygoid a joint is formed. This is why in the adults of these 
birds the pterygoid seems not to reach the prevomer, whereas morphologically, in 
fact, it does or almost does reach it. The hemipterygoid in various Carinates is 
shown in Plates 8 and 9 for comparison with the conditions in Ratites. 

For those, if there be any, who still believe in the theory of recapitulation, it 
would no doubt be tempting to say that the neognathous palate " recapitulates " in 
its development the condition of the palaeognathous palate which would therefore 
be ancestral. But in view of the overwhelming evidence that the Ratites are 
secondarily descended from flying birds, the fact that the Ratites already show 



THE EVOLUTION OF RATITES 67 

neoteny in two other features, the plumage and the bones of the skull, and the 
probability, from A. Kleinschmidt's (1951) reconstruction, that the palate of 
Archaeopteryx was neognathous (schizognathous) , it is impossible to believe that in 
their palates the Ratites are primitive. The palaeognathous type of palate must 
therefore be neotenous. This means a complete reversal of the hitherto generally 
held view of the palate of birds and necessitates the conclusion that the so-called 
neognathous palate is primitive. 

The primitive nature of the neognathous palate in birds is probably connected 
with the phenomenon of kinetism. J. Versluys (1910) has shown that the mesokin- 
etic condition in Carinates, where the quadrate and pterygoid bones are capable of a 
certain amount of movement and sliding, and there is a joint between the pterygoid 
and palatine whereby the upper jaw can be moved on a hinge at the level of the 
lacrimalbones and raised relatively to the brain-case, is only intelligible if the birds 
were evolved from reptiles in which a similar though less extensive power of movement 
was possible : the condition which he has called met akinetic. According to him 
(1910 : 244) even Archaeopteryx had a kinetic skull capable of movement, although it 
still possesses a preorbital bar separating the preorbital fossa from the orbit, and a 
suborbital bar. But in the Ratites the power of movement has been reduced ; the 
quadrate has a broad connexion with the pterygoid, the latter has equally broad 
connections with the palatines and the pre vomer, and there is no movable joint 
between the pterygoid and the hemipterygoid because these two elements have not 
become separated. It must be concluded therefore that with the loss of flight, 
general increase in size, and acquisition of different feeding habits, the Ratites have 
lost the Carinates' power of movement of the upper jaw, by retaining the juvenile 
condition of the palate before any joint is formed. I am greatly indebted to Dr. 
W. C. Osman Hill for informing me that even in the kiwi, which is the smallest of 
the Ratites, there is no mobile joint at the base of the upper jaw ; and that in 
the cassowary the only very slight mobility in the upper jaw is at a point far 
forward, just behind the nostrils. 

Further, there is a curious point in the distribution of the palaeognathous type of 
palate among the birds. It is found not only in Ratites, but also in the tinamus, 
which are Carinates with a well-developed keel on the sternum and good power of 
flight. This fact in itself is sufficient to indicate that the Ratites have lost the power 
of flight, for it could hardly be contended that the tinamus have evolved flight from 
a flightless Ratite condition. 

As for the argument that the absence of any rudiment of the clavicle in the ostrich 
implies that it was evolved from ancestors which lacked the clavicle (and, by implic- 
ation, could not fly), it is another example of the fallacies to which the theory of 
recapitulation leads by its assertion that early embryonic stages of development 
must represent early ancestral stages in evolution. Modern birds lack even the 
rudiments of teeth, but teeth are present in Archaeopteryx, Hesperornis, and 
Ichthyornis. The absence of tooth rudiments in modern birds no more excludes 
Archaeopteryx from their ancestry than the absence of limb-rudiments in snakes 
indicates that their ancestors were limbless. 

Finally, the embryonic development of the emu, studied by H. Steiner (1936) 



68 THE EVOLUTION OF RATITES 

and H. Lutz (1942), shows that the structure and organization of the Ratite embryo 
is so similar to the Carinate that it can only be interpreted on the view that Ratites 
have evolved from flying birds. 

CONCLUSIONS 

On all these grounds, therefore, there can be no doubt that Owen was correct in 
regarding the Ratites as birds which have " abrogated " the power of flight. It is 
possible to go further and to say that they have degenerated from a Carinate 
condition. Whether the Ratites represent a natural group or whether they are 
an assemblage of forms which have independently followed parallel lines of evolution 
consequent on the loss of flight is a further problem for ornithologists to solve. 

In view of the incontrovertible evidence from the structure of the wing, the 
pygostyle, and the cerebellum, that the Ratites have degenerated from flying birds, 
any attempt to explain the persistent juvenile characters of the Ratites (nestling- 
down, skull-sutures, and palate) as phylogenetically primitive is doomed to failure ; 
and the Ratites must be regarded as providing one of the most telling exposures of 
the fallacy of the theory of recapitulation. 

I am glad to acknowledge the help of my colleagues in the Bird Room of the 
British Museum (Natural History), Mr. J. D. Macdonald and Miss P. A. Lawford, of 
Dr. W. E. Swinton of the Department of Geology, and of Mr. J. V. Brown, Senior 
Photographer. 

SUMMARY 

Now that the anatomy of Archaeopteryx is adequately known, it is possible to make 
a rigorous analysis of the characters of the Ratites in the light of the conditions 
shown by primitive birds. The structure of the wing, tail, and brain in Carinates 
shows advances on Archaeopteryx which are undoubtedly adaptations to flight. The 
presence of the same features in Ratites proves that they are descended from flying 
birds. The condition of the plumage, the sutures between the bones of the skull, 
and the disposition of the bones of the palate in Ratites, all show secondary retention 
of characters which are juvenile in Carinates, and are evidence of neoteny in the 
Ratites. 

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McDowell, S. 1948. The bony palate of birds. Part I : The Palaeognathae. The Auk, 

65 : pp. 520-549. 
Owen, R. 1866. On the Anatomy of Vertebrates. II. Birds and Mammals. London, i-viii, 

586 pp. 
Parker, T. J. 1892. Observations on the anatomy and development of Apteryx. Philos. 

Trans. B. 182, 25-134. 
Parker, W. K. 1875. On the morphopgy of the skull in the Woodpeckers (Picidae) and 

Wrynecks (Yungidae). Trans. Linn. Soc. London, Zool., 1 : 1-22. 

1876. On the structure and development of the bird's skull. Trans. Linn. Soc. London, 

Zool., 1 : 99-154. 

1877. On Aegithognathous birds (Part I). Trans. Zool. Soc. London, 9 : 289-352. 

1879. On the skull of Aegithognathous birds (Part II). Trans. Zool. Soc. London, 

10 : 251-314. 

Pycraft, W. P. 1900. On the morphology and phylogeny of the Palaeognathae (Ratitae 

and Crypturi) and Neognathae (Carinatae). Trans. Zool. Soc. London, 15 : 149-290. 
— — 1900. On the palate of Caprimulgidae. Bull. Brit. orn. CI. xi : 12-13. 

1 90 1. Some of the points in the morphology of the palate of the Neognathae. Journ. 

Linn. Soc. London, Zool., 28 : 343-357. 

Simpson, G. G. 1946. Fossil Penguins. Bull. Amer. Mus. nat. Hist. 87 : 7-99. 



70 THE EVOLUTION OF RATITES 

Steiner, H. 1918. Das Problem der Diastataxie des Vogelfliigels. Jena Z. Naturw. : 

55 : 221-246. 

1936. Ueber die aussere Gestaltung ernes fimfzehntagigen Embryos des Emus. Rev. 

Suisse Zool. Geneve, 43 : 543-550. 

1938. Der " Archaeopteryx " Schwanz der Vogelembryonen. Vjschr. naturf. Ges. 

Zurich, 83, Beiblatt : 279-300. 
1949- Zur Frage der ehemaligen Flugfahigkeit der Ratiten. Rev. Suisse Zool. Geneve, 

56 : 364-370. 

1956. Die taxonomische und phylogenetische Bedeutung der Diastataxie des Vogel- 
fliigels. J . Orn., Lpz., 97 : 1-20. 

Stresemann, E. 1927-1934. Sauropsida ; Aves, etc., xi, 899 pp. in Kuekenthal, Handbuch 
der Zoologie, 7 : Halfte II. 

Tucker, B. W. 1938a. Functional evolutionary morphology : the origin of birds. Evolu- 
tion, Essays on Aspects of Evolutionary Biology : edited by G. R. de Beer, Oxford, 321-336. 

i93 8 &- Some observations on Dr. Lowe's theory of the relationship of the Struthiones 

to the Dinosaurs and to other birds. Proc. 8th Int. orn. Congr. Oxford, 1934 : 222-224. 

Versluys, J. 1910. Streptostylie bei Dinosaurien, nebst Bemerkungen iiber die Verwand- 
schaft der Vogel und Dinosaurier. Zool. Jahrb. Jena (Abt. Anat. & Ontogenie d. Tiere) : 
30 : 175-260. 

Wray, R. S. On some points in the Morphology of the Wings of Birds. Proc. Zool. Soc. 
London : 343~357- 

Young, J. Z. 1950. The Life of Vertebrates. Oxford. 



EXPLANATION OF PLATES 

PLATE 5 

(1) Right side view of the pygostyle in a Carinate, Leptoptilos crumeniferus (Marabou Stork) and 
(2), in a Ratite, Stvuthio camelus, showing the similarity of structure, (ap), the anterior portion showing 
the elements of a distinct vertebra ; (pp) posterior portion composed of fused vertebrae. 

PLATE 6 

(1) The brain as seen in right-side view in Archaeopteryx lithographica. (2) sagittal section of the 
brain in a Carinate, Tringa ocrophus (Green Sandpiper), and (3) in a Ratitie, Rhea americana. (ce), 
cerebellum ; (ch), cerebral hemisphere ; (ol), optic lobes. 

PLATE 7 

The arrangement of the feathers on the wing of a young Ratite, Rhea americana, showing the differen- 
tiation between primary and secondary remiges, bastard wing, and wing-coverts. 

PLATE 8 

(1) Ventral view of the structure of the palate in the Carinate Pygocelis papus (Gentoo penguin) 
nestling ; (2) in the Carinate Anthropoides paradisea (blue crane) ; and (3) in the Ratite Dromiceus 
novae-hollandiae (emu), (hpt), hemipterygoid ; (pa), palatine ; (pt) pterygoid ; (pr), prevomer ; (qu), 
quadrate. 

PLATE 9 

(1) The structure of the palate as seen in left-side view in Corvus frugilegus (rook) young ; (2), Mega- 
laema virens (Himalayan barbet). (hpt), hemipterygoid ; (ju), jugal ; (pa), palatine ; (pt), pterygoid ; 
(pr), prevomer ; (qu), quadrate. 






Bull. B.M. {N.H.) Zool. 4, 2 



PLATE 5 




PLATE 5 
(1) Right side view of the pygostyle in a Carinate, Leptoptilos crumeniferus (Marabou Stork), 
and (2), in a Ratite, Struthio camelus, showing the similarity of structure, (ap), the anterior 
portion showing the elements of a distinct vertebra ; (pp) posterior portion composed of fused 
vertebrae. 



Bull. BM. {N.H.) Zool. 4, 2 



PLATE 6 



ce 0/ ch 



ce ch 




PLATE 6 

(1) The brain as seen in right-side view in Archaeopteryx lithographica. (2) sagittal section 
of the brain in a Carinate, Tringa ocrophus (Green Sandpiper), and (3) in a Ratite, Rhea ameri- 
cana. (ce), cerebellum ; (ch), cerebral hemisphere ; (ol), optic lobes. 



Bull. B.M. {N.H.) ZooL 4, 2 



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PLATE 8- 




pv pa hpt pv pa hpt pt pt ' 




PLATE 8 

(1) Ventral view of the structure of the palate in the Carinate Pygocelis papas (Gentoo pen- 
guin) nestling ; (2) in the Carinate Anthropoides paradisea (blue crane) ; and (3) in the Ratite 
Dromiceus novae-hollandiae (emu), (hpt), hemipterygoid ; (pa), palatine ; (pt) pterygoid ; 
(pv), prevomer ; (qu), quadrate. 



Bull. B.M. (N.H.) Zool. 4, 2 



PLATE 9 





PLATE .9 

(1) The structure of the palate as seen in left-side view in Corvus frugilegus (rook) young 
(2), Megalaema virens (Himalayan barbet). (hpt), hemipterygoid ; (ju), jugal ; (pa), palatine 
(pt), pterygoid ; (pr), prevomer ; (qu), quadrate. 



PRINTED IN GREAT BRITAIN BY 
ADLARD AND SON, LIMITED, 
BARTHOLOMEW PRESS, DORKING 



2 rmrim STUDIES ON THE 
TRICHIUROID FISHES— 3 



A PRELIMINARY REVISION OF THE 
FAMILY TRICHIURIDAE 



DENYS W. TUCKER 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 4 No. 3 

LONDON: 1956 



STUDIES ON THE TRICHIUROID FISHES— 3 

A PRELIMINARY REVISION OF THE FAMILY 

TRICHIURIDAE 




BY 

DENYS W. TUCKER 



Pp. 73-130 ; PI. 10 ; 23 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 4 No. 3 

LONDON: 1956 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 4, No. 3 of the Zoological series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued August, 1956 Price One Pound 



STUDIES ON THE TRICHIUROID FISHES— 3 

A PRELIMINARY REVISION OF THE FAMILY 

TRICHIURIDAE 

By DENYS W. TUCKER 

CONTENTS 

Page 
Introduction ........... 74 

The Characters of the Family Trichiuridae ..... 74 

A Short Key to the Subfamilies and Genera of the Family Trichi- 
uridae ............ 77 

Systematic Review : 

Subfamily Aphanopodinae ........ 77 

Genus Diplospinus ......... 78 

,, Aphanopus ......... 81 

,, Benthodesmus . . ... . . . .85 

Subfamily Lepidopodinae ........ 89 

Genus Lepidopus ......... 90 

,, Evoxymetopon ........ 97 

,, Eupleurogv animus . . . . . . . .102 

,, Assurger ......... 106 

Tentoriceps . . . . . . . . .110 

Subfamily Trichiurinae . . . . . . . • .112 

Genus Trichiurus . . . . . . . • • 113 

Leptur acanthus . . . . . . . .119 

The Origin, Evolution and Classification of the Trichiuridae : 

Summary of earlier work . . . . . . • .120 

Nesiarchus-Diplospinus : the Gempylid-Trichiurid bridge . . .122 

Evolutionary trends in the Trichiuridae . . . . . .125 

Classification of the Trichiuridae . . . . . . .128 

References ........••• 129 

SYNOPSIS 

This paper presents a working classification of the Trichiuridae, based on a consideration of 
the literature of the family and examination of selected material, which has been prepared as 
a prelude to Reports on the " Dana " collections of Trichiuridae and Gempylidae. Three 
subfamilies are recognized : Aphanopodinae (genera Diplospinus, Aphanopus, Benthodesmus) ; 
Lepidopodinae (genera Lepidopus, Evoxymetopon, Eupleurogrammus , Assurger, Tentoriceps) and 
Trichiurinae (genera Trichiurus, Leptur acanthus) . Keys, diagnoses and synonymies are presented 
and figures given of all species recognized. Diplospinus is considered to be the most primitive 
genus extant and grounds are given for relating it to the Gempylid Nesiarchus. The origin and 
radiation of the Trichiuridae are discussed. 

1 The previous papers in this series were : (1) The fishes of the genus Benthodesmus (Family Tri- 
chiuridae), Proc. zool. Soc. Lond. 123 : 171-197, 3 pis., 5 text-figs. (1953). (2) Benthodesmus tenuis 
(Gunther) collected by the Expedition Oceanographique Beige dans les eaux cotieres de l'Atlantique 
Sud (1948-1949), with additional notes on the genus Benthodesmus, Bull. Inst. roy. Sci. nat. Belg. 31, 
No. 64 : 1-26, 1 pi., 11 text-figs. (1955). 

zool. 4, 3. 6 



74 



THE FAMILY TRICHIURIDAE 



INTRODUCTION 



As I contemplated the mass of material which resulted from my rash acceptance of 
Dr. Anton Fr. Bruun's invitation to write reports on the young Trichiuroid fishes 
collected by the "Dana " Expeditions, I realized the urgent need of some preliminary 
working classification with which to regulate the chaos that must ensue once these 
many thousands of specimens were released, like so many djinns, from their tubes 
and bottles. 

The problem of the Gempylidae was immediately relieved by Matsubara and Iwai 
(1952) and by Mrs. Marion Grey (1953), but the case of the Trichiuridae remained 
desperate. There has been no comprehensive revision of this family since the end of 
the nineteenth century. The earlier synopses of Gunther (i860), Gill (1863) and 
Goode and Bean (1895) are no longer adequate accounts even of the genera which 
they describe and, moreover, contain no attempt at a phyletic classification since 
they date from a period before the planting of family trees became fashionable. 
Later workers have had varying success in distinguishing the genera and species 
of limited regions. In this century a few new species and genera have been proposed, 
two of the latter without any of the inhibitions consequent upon an interest in the 
family or the possession of study-material. 

The present draft revision assigns a place to every nominal genus and species 
and gives, as a minimum, the reference for the first publication of every name and 
name-combination, together with selected items from the remaining literature. It 
gives diagnoses and a phyletic classification of all sub-families, genera and species 
recognized and argues the case for synonymies with whatever detail the individual 
circumstances may immediately demand. Except for Evoxymetopon, Assurger and 
Tentoriceps (of which material or new published descriptions would be greatly 
appreciated), material of all genera and species has been examined, including a 
substantial number of type specimens. 

The author of any " preliminary " contribution should justify his title. The amount 
of labour involved in preparing the present MS as a working tool has shown the need 
of such a tool and of certain small but critical contributions to the understanding 
of the Trichiuridae which those possessing rarer material may make. It will be some 
considerable time before the final "Dana " Reports on the Trichiuridae and Gempy- 
lidae can be completed and so, faute-de-mieux , a preliminary account appears likely 
to be useful, even though some of its conclusions may be subject to second thoughts. 

I wish to express my thanks to Messrs. P. E. Purves and A. C. Wheeler of the 
British Museum (Natural History) for numerous radiographs which have been of 
very great assistance in this work. 

THE CHARACTERS OF THE FAMILY TRICHIURIDAE 

Regan (1909) allies the Trichiuridae with the Gempylidae as the Trichiuriformes, 
forming the first division of his suborder Scombroidei of the order Percomorphi. 
He characterises the Trichiuriformes as having : — 

" Caudal fin-rays not deeply forked at the base, the hypural in great part 



THE FAMILY TRICHIURIDAE 75 

exposed. Praemaxillaries beak-like, free from the nasals ; mouth toothed, 
with lateral cJeft ; strong anterior canines. Epiotics separated by supra- 
occipital. Gill-membranes free from the isthmus. Pectoral fins placed low." 

With this diagnosis I have no present disagreement save to comment that hypurals 
are sometimes absent and to prefer the use of " fangs " or " caniniform teeth " 
rather than " canines ' for fish teeth ; the term " canine " is best restricted to 
certain reptiles and to the mammals, in which it is defined, not by form but by 
position and homology, as " the most anterior tooth of the maxilla, situated on or 
immediately behind the premaxillo-maxillary suture ... or the tooth in the lower 
jaw which bites in front of the upper canine ". 

Regan's diagnosis of the family Trichiuridae follows : 

" Body very elongate, strongly compressed ; maxillary sheathed by the 
praeorbital ; spinous dorsal, if distinct, not longer than the soft 1 ; anal with 
numerous short spines 2 ; pelvic fins reduced to a pair of scale-like appendages 
or absent 3 ; caudal small or absent. Dorsal and anal rays corresponding to 
the vertebrae 4 , each interneural or interhaemal attached to a neural or haemal 
spine ; pelvic bones, if present, united to form a slender spicular bone connected 
with the cleithra by a long ligament 5 . Vertebrae numerous, 100(43 + 57) to 
I 59(39 + I2 °) or more 6 ; ribs feeble, sessile." 

This description is evidently based primarily upon examinations of Lepidopus, 
Aphanopus and Trichiurus and requires several modifications and qualifications : 

(1) The spinous dorsal is always distinct ; it is longer than the soft in Diplospinus 
(discovered since Regan's time) and very slightly longer than the soft in occasional 
specimens of Aphanopus. 

(2) Some, if not all, of the anal rays are split, soft and support a fin-membrane 
(Diplospinus, Aphanopus, Benthodesmus, Lepidopus, Evoxymetopon, Assurger) ; 
in Trichiurus, Leptur acanthus and Eupleurogrammus, however, the anal rays are much 
reduced spinules or entirely absent. At the origin of the anal fin, moreover, immediately 
behind the vent, are two spines (represented by the notation i -f- I throughout the 
present paper) ; of these the anterior is a minute spinule while the second may be 
variously enlarged as a leaf -like or keeled scute, or as a stout spine. 

(3) The pelvic fins in some genera (Diplospinus, Aphanopus, Benthodesmus, 
Lepidopus) and probably in all in which they are present, consist each of a scale-like 
spine and one rudimentary soft ray, the latter newly noticed. 

(4) The dorsal spines and their basals and interneurals always correspond to the 
trunk vertebrae ; the dorsal soft rays may be twice as numerous as the adjacent 
vertebrae (Diplospinus), slightly more numerous (Aphanopus, Benthodesmus) or as 
numerous (remaining genera) . 

(5) The pelvic bones form an imperfectly fused, fenestrated structure which is not 
always elongated. 

(6) The vertebrae range from 34 + 24 = 58 (Diplospinus) to 53 + I0 3 = x 56 
(Benthodesmus simonyi) or 41 + 151 = 192 (Eupleurogrammus muticus). 



7 6 



THE FAMILY TRICHIURIDAE 





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THE FAMILY TRICHIURIDAE 77 

A SHORT KEY TO THE SUBFAMILIES AND GENERA 
OF THE FAMILY TRICHIURIDAE 

Frontal ridges not elevated, no sagittal crest. Profile of head rising very gently from 

snout tip to dorsal (cf. Text-fig. 1) . . . . . Aphanopodinae (p. 77) 

D. 72-73. Spinous dorsal base twice as long as soft . . . Diplospinus (p. 78) 

D. 8 2-8 7. See " Lepidopus xantusi " (Lepidopodinae) 

D. 91-95. Spinous and soft dorsal bases sub-equal .... Aphanopus (p. 81) 

D.120+ Spinous dorsal base half as long as soft . . . . Benthodesmus (p. 85) 

Posterior confluence of frontal ridges elevated, forming a prominent sagittal crest at the 

nape, which may or may not be continued forward as a ridge-like elevation of the 

ethmo-frontal region (cf. Text-figs. 2, 3 and 4). 

Ventral fins present. Lateral line descending gently from the shoulder and median or 

sub-median along the body, i.e. distance from lateral line to ventral profile at 

anus much more than half distance from lateral line to dorsal. Lower hind 

margin of operculum convex ...... Lepidopodinae (p. 89) 

Sagittal crest confined to nape. Interorbital concave. Caudal present 

Lepidopus (p. 90) 
Sagittal crest continuous from snout tip to dorsal. Interorbital convex 
Caudal present 

D. 87-93. Body depth 12-13 m length ..... Evoxymetopon (p. 97) 

D.120. Body-depth 20-28 in length . ... . . . Assurger (p. 106) 

Caudal absent. 

Body depth 14-18 in length . . . . Eupleurogrammus (p. 102) 

Body depth 20-24 in length ...... Tentoriceps (p. no) 

Ventral fins absent. Lateral line descending steeply from the shoulder and running 
near the ventral profile of the body, i.e. distance from lateral line to ventral profile 
at anus less than half distance from lateral line to dorsal. Lower hind margin of 
operculum more or less concave. Caudal always absent (cf. Text-fig. 4) 

Trichiurinae (p. 112) 
Post-anal scute small, less than the pupil. Soft anal rays not breaking through 

skin. Eye large, 5 . 0-7 . o in head ..... Trichiurus (p. 113) 

Post-anal scute large, half the eye-diameter. Soft anal rays pungent spinules, 

breaking ventral profile. Eye small, 6.7-10.0 in head . . Leptut 'acanthus (p. 119) 

SYSTEMATIC REVIEW 

Subfamily APHANOPODINAE Gill 

Aphanopodinae Gill, 1863, Pvoc. Acad. nat. Sci. Philad. 1863 : 225. 
Type genus Aphanopus Lowe. 

Genera now recognised. — Aphanopus Lowe ; Benthodesmus Goode & Bean ; 
Diplospinus Maul. 

Diagnosis : 

A. Snout gently sloping ; orbits entering upper profile of head ; frontal ridges 
only slightly elevated, not contributing to a sagittal crest. 

B. A stout, conical, cartilaginous protuberance at the mandibular symphysis ; 
another, much smaller, at the tip of the snout. 

C. Lower hind margin of operculum markedly convex. 



78 THE FAMILY TRICHIURIDAE 

D. Teeth of main series with double barbs (Diplospinus) or entirely without 
barbs (Aphanopus, Benthodesmus) . 

E. Teeth on palatines in a linear series. (In Aphanopus only 1-2 posterior 
rudiments of the series present.) 

F. Lateral line descending gently from the shoulder and running in a median 
or sub-median position along the body, i.e. distance between lateral line 
and ventral profile much more than half distance between lateral line and 
dorsal. 

G. Spinous dorsal fin long, with 32-46 rays. Spinous and soft dorsals partly 
divided by a slight notch. 

H. Soft dorsal rays slightly more numerous than adjacent caudal vertebrae, or 
up to twice as many. Basal and interneural elements intercalated among 
the main series and unrelated to neural spines of vertebrae. 

I. Spinous anal i + I ; anterior soft anal rays weak but (except in Benthodesmus 
simony i) an external fin is continuous in some form or other from the vent 
nearly to the caudal ; the properly developed fin may extend the whole 
length or be confined to the posterior 20-25 rays. 

J. Terminations of dorsal and anal fins sub-opposite. 

K. Caudal fin always present ; small, normal, forked. 

L. Ventral fins always present (though reduced to internal rudiments in adult 
Aphanopus), composed each of a scale-like spine and one soft ray ; in the 
adult fish inserted not more than 2-3 mm. before/behind anterior /posterior 
perpendiculars through the ends of the pectoral base. 

M. Pyloric caeca few (6-9) ; (not verified in Diplospinus). 

Osteological Literature 

Giinther, i860, Cat. Fish. B.M. 2 : 342-344 (desc. osteology Aphanopus). 
Tucker, 1953, Proc. zool. Soc. Lond. 123 : 196-197, pis. 2-3 (figs, osteology of paired fins 
and anal fin of Aphanopus & Benthodesmus). 

1955, Bull. Mus. Hist. nat. Belg . 31, No. 64 : 1-26 (figs, osteology of pelvic and anal fin 

of Benthodesmus) . 

Literature on young stages 

Maul, 1948, Bol. Mus. Funchal No. 3, Art 6 : 42, fig. 17 (young Diplospinus). 
Tucker, 1953, °P- c ^. : 187 (figs, young Aphanopus and Benthodesmus). 



Genus DIPLOSPINUS Maul 

Diplospinus Maul, 1948, Bol. Mus. Funchal No. 3, Art. 6 : 42. 

Type species Diplospinus multisiriatus Maul. Monotypic. 

Synonyms 

Lepidopus (non Gouan 1770) (part) Brauer, 1906. 

Benthodesmus (non Goode & Bean 1882) (part) Goode & Bean, 1895 ; Fowler, 1938. (Refs. 
below.) 



THE FAMILY TRICHIURIDAE 



79 



Diagnosis : 

(i) Body elongate, head length 6-6-6-9 in standard length 125-203 mm., 
body depth 18-5 in S.L. 

(2) Vent exactly in middle of S.L. 

(3) Vertebrae 34 + 24 = 58. (Corresponds to 36 + 22 = 58 in convention 
used for Benthodesmus.) 

(4) Spinous dorsal base twice as long as soft dorsal base. 

(5) Dorsal spines 32-33 ; dorsal soft rays 40. 

(6) Dorsal soft rays about twice as numerous as adjacent caudal vertebrae, so 
that alternate interneural elements do not articulate with neural spines. 

(7) Anal spines i + I, the former half the length of the latter in young stages ; 
condition in the adult unknown ; i is linear ; I is dagger-shaped and V-shaped 
in transverse section. 

(8) Anal spines i and I articulate close together on a common basal, which is 
not enlarged or specially modified and which, except that it does not quite 
touch the corresponding haemal arch, does not show any difference in the 
size and relations of the interhaemal process from those which follow it. 
(Condition similar to Lepidopus.) 

(9) A complete external anal fin supported by 31 split but unbranched rays 
extending from the spinous anal nearly to the caudal. The soft rays and 
their basal elements are about twice as numerous as the adjacent caudal 
vertebrae, so that alternate basals have interhaemal processes which are 
unrelated to haemal arches. 

(10) Ventral fins inserted on perpendicular through anterior end of pectoral 
fin-base. 

(11) Ventral fin I-i ; a narrow scale-like spine and an external split ray twice 
as long. 

(12) All principal teeth of the premaxillary and dentary series are strongly 
barbed (arrowhead-shaped), with thickened enamel caps. 

(13) Palatine teeth in a linear series, exposed. 

(14) Principal teeth on first gill-arch numerous. 

(15) Long intermuscular (pleurals and epipleurals) bones present, extending 
throughout trunk. 

(16) Melanophores distributed in parallel and narrow longitudinal rows along 
the body. 

One species, Diplospinus multistriatus Maul, Atlantic and Indo-Pacific. 

Diplospinus multistriatus Maul, 

(Text-fig. 5) 
Benthodesmus atlanticus (part) Goode & Bean, 1895, Oceanic Ichthyology : 206 (the two small 

specimens mentioned, fide Dr. Carl L. Hubbs, in litt.). 
Benthodesmus benjamini (part) Fowler, 1938, Proc. U.S. Nat. Mus. 85 : 45 (certain of the para- 
types, fide Dr. Carl L. Hubbs, in litt.). 
? Lepidopus gracilis Brauer, 1906, Wiss. Ergeb. " Valdivia " 15 : 291, Taf. XII, fig. 1 (not fig. 5 
as erroneously stated in the text nor fig. 3 as stated in the legend to the plate). 

Holotype in the Berlin Museum ? Type locality West coast of S, Africa, St. 82, 
21 53' S., 6° 58' 6" E. 



8o 



THE FAMILY TRICHIURIDAE 



3 



THE FAMILY TRICHIURIDAE 81 

Diplospinus multistriatus Maul, 1948, Bol. Mus. Funchal, No. 3, Art 6 : 42, fig. 17. 
Holotype Museu Municipal do Funchal No. 3063. Type locality Madeira. 
Paratypes Museu Municipal do Funchal Nos. 3064-5, 3067-9. 
Paratype British Museum (Natural History) No. 1953.10.28.1. (Formerly 3066.) 

Certain discrepancies will be noticed between the generic diagnosis given above 
and the otherwise accurate description and figure by Maul (1948) ; the corrected 
observations have been made on the paratype kindly presented by Mr. G. E. Maul. 
Each ventral fin includes a soft ray in addition to the spine ; there is a single row 
of about a dozen teeth on each palatine (" no teeth on vomer or palatines ") ; there 
are traces of an apparent and highly probable lateral line (" no lateral line ") though 
the present specimen is completely skinned ; certain of the premaxillary fangs are 
represented by replacement teeth (" depressible teeth "). The number of branchio- 
stegal rays is 7, as in other Trichiurids. The number of pyloric caeca cannot be 
determined owing to destruction of the thoracic region. There is a deep notch on the 
hinder margin of the opercular, as already observed by Maul, and this character 
proves to be rather important since it is confined to Diplospinus, the most primitive 
recent Trichiurid and to Nesiarchus, the nearest-related Gempylid (see p. 124). 

Since Brauer's (1906) figure of Lepidopus gracilis bears the magnification 2/1 we 
may deduce a S.L. of 68 mm., i.e. about one-third the length of the type series of 
Diplospinus multistriatus. The head is 4-8 and the height 14-4 in the length ; the 
eye goes 5 times in the head, and the ventral and anal spines are proportionately 
longer than in the types. All these differences are in the directions to be expected in 
a younger fish. The counts of D.65-67 and A.27 are slightly low, but not outside 
the probable range of variation or error. However, the eye is shown about a quarter 
its diameter below the dorsal profile of the head, the origin of the dorsal fin is a little 
retarded and the insertion of the ventral fins is below the posterior rather than the 
anterior end of the pectoral base (" Bauchflosse kurz hinter der Vertikale der 
Brustflosse "). These discrepancies must await a satisfactory explanation, which is 
likely to result in Diplospinus gracilis (Brauer) becoming the definitive name of the 
present species. 

Genus APHANOPUS Lowe 
Aphanopus Lowe, 1839, Proc. zool. Soc. Lond. 7 : 79. 

Type species Aphanopus carbo Lowe. Monotypic. 

Synonyms 

Lepidopus (non Gouan, 1770) Sim, 1898 ;Dons, 192 1. (Refs. below). 

Diagnosis : 

(1) Body elongate, head length 5-68-4-92 in standard length 102-1036 mm., 
body depth 21-7-11-23 in same. 

(2) Tail 48-49% of standard length. 

(3) Vertebrae 42-44 + 55-56 = 98-99. 

(4) Spinous and soft dorsal bases sub-equal, differing by at most ± 3 % of S.L. 

(5) Dorsal spines 38-41 ; dorsal soft rays 53~56 ; aggregate 91-95. 

(6) Dorsal soft rays practically corresponding with adjacent caudal vertebrae. 



82 



THE FAMILY TRICHIURIDAE 




Oh 

I"? 

a; 

Oh 

J a, 

S § 

N rrl 

s ^ 

M 

H 



I 



THE FAMILY TRICHIURIDAE 83 

There are very few intercalated interneural elements which are unrelated 
to neural spines and these occur usually towards the beginning and end 
of the fin. 

(7) Anal spines i + I, the former about 1 /5 the length of the latter in the young 
stages but becoming disproportionately smaller in the adult, in which i 
becomes a minute sharp spinule, usually concealed beneath the skin and I 
is a stout dagger-shaped spine, triangular in cross-section. 

(8) Anal spine i articulates a short distance in advance of I. Their common 
basal element is a complex, greatly enlarged and strengthened to accommo- 
date the hyper trophied I, and representing four or more fused elements. The 
compound interhaemal process is stout and does not touch the adjacent 
haemal arch. The horizontally directed component of the compound 
basal element occupies the length of three vertebral centra and the presumed 
anterior migration of the corresponding interhaemal processes leaves a 
space above it. 

(9) A complete external anal fin of 44-48 split but unbranched rays extending 
from the spinous anal nearly to the caudal. The anterior rays are very 
weak and the functional fin is effectively confined to the posterior 20-25 
rays. The internal supporting skeleton is quite regular ; there is a precise 
correspondence between rays, basal elements and caudal vertebrae, with 
a close association between interhaemal processes and haemal arches. 

(10) Ventral fins inserted immediately before perpendicular through anterior 
end of pectoral fin-base. External fins present only in the juvenile ; 
fins and girdle reduced to an internal rudiment in the adult. 

(11) Ventral I-i in the juvenile only ; a narrow spine and an external split ray 
initially about 3 times as long. 

(12) Principal teeth of the premaxillary and dentary series without barbs : 
if these are sometimes present on the premaxillary fangs they are usually 
barely perceptible and confined to the hinder edges, without enamel 
thickening. The marginal teeth of the jaws are stout, triangular and have 
microscopically-serrated edges. 

(13) Palatine teeth reduced to 1-2 minute rudiments at hinder end of bone, 
very much concealed. 

(14) Principal teeth on first gill-arch very numerous. 

(15) Intermuscular bones (pleurals and epipleurals) weaker than in Diplospinus. 

(16) Pigmentation uniform, dense ; fish uniform black when dead. Living fish 
coppery with iridescent reflexions. 

One species, Aphanopus carlo Lowe, N. Atlantic and Gulf of Aden. 

Aphanopus carbo Lowe 
(Text-figs. 6 & 7). 
Aphanopus carbo Lowe, 1839, Proc. zool. Soc. Lond. 7 : 79. 

Holotype B.M. (N.H.) No. 1851 . 1 1 . 29 . 6. Type locality Madeira. 
Aphanopus minor Collett, 1886, Chr. Vid.-Selsk. Fork. 1886 No. 19 : 1, fig. 1. 

Holotype in Universitetets Zoologiske Museum, Oslo. Type locality Denmark Strait, 
E. of Greenland, 65 N., 31 W. 







Table I. 






Holotype. 


Vertebrae. 


Dorsal. 


A. 


carbo . 


42+56 


XXXVIII, 56 


A. 


schmidti 


42+56 


XXXVIII, 55 


A. 


minor . 


44+ ? 


XLI, ? 


A. 


microphthalmus 


44+55 


XLI, 54 



84 THE FAMILY TRICHI URID AE 

Lepidopus caudatus [non Euphrasen, 1788) Sim, 1898, Ann. Scot. nat. Hist. 1898 : 53. 
Aphanopus schmidti Saemundsson, 1907, Vid. Medd. naturh. Foren. Kbh. 59 : 22, PL 1. 

Holotype in Natturugripasafnid, Reykjavik. 

Paratype B.M. (N.H.) No. 1925.7.23.4. Type locality Vestmann Is., S.W. of Iceland. 
Lepidopus atlanticus {non Goode & Bean, 1895) Dons, 1921, Tromso Mus. Aarsh. 43, No. 6 : 10, 
fig. 1. 

(Identification corrected to Aphanopus schmidti by Soot-Ryen, 1936, Nytt. Mag. Naturv. 
76 : 237.) 
Aphanopus microphthalmus Norman, 1939, Sci. Rep. John Murray Exped. 7 No. 1:71, fig. 25. 

Holotype B.M. (N.H.) No. 1939.5.24. 1322. Type locality Gulf of Aden. 
Aphanopus acus Maul, 1948, Bol. Mus. Funchal No. 3, Art. 6 : 47, fig. 18. 

Holotype in Museu Municipal do Funchal. Type locality Madeira. (Withdrawn as 
young A. carbo by Maul, 1949, Bol. Mus. Funchal, No. 4, Art. 10 : 21.) 
non Aphanopus simonyi Steindachner, 1891. (See under Benthodesmus simonyi.) 
non Aphanopus carbo Norman, 1937. (Mediterranean records based on confusion with Lepidopus 

caudatus, q.v.) 



Anal. 

i+I+48 
i+I+46 
i+I+ ? 
i+I+45 

In the type of A. minor the tail has been broken off a short distance behind the vent and has 
subsequently healed over with some slight re-orientation of the soft dorsal and anal rays 
remaining. The remnant includes 25 caudal vertebrae, 28 soft dorsal rays, 21 anal elements. 

Through the kindness of Dr. C. Stop-Bowitz (Oslo), Dr. Finnur Gudmundsson 
(Reykjavik) and Mr. G. E. Maul (Funchal) I have been able to examine the types 
of all the nominal species of Aphanopus and, by comparing these with a series of 
some thirty specimens from the type locality and as many more from the North 
Atlantic, to decide that they represent only one species, A. carbo Lowe. 

A. acus Maul is a juvenile A. carbo and has already been adequately dealt with by 
Maul (1949). Meristic counts for the other nominal species are given in Table I. 
Ranges of vertebral counts for the long series are not yet available, but the variations 
now tabulated are small and well within the limits of those found in Benthodesmus 
tenuis (p. 88). Fin-ray counts on eighteen Madeiran specimens give ranges 
D.XXXVIII-XL, 53-55 (aggregate 91-95) ; A. i + I + 44-4$. 

The validity of A. schmidti has been much debated, Saemundsson pro, Grieg 
and others con. The arguments will be dealt with in detail elsewhere ; for the present 
it is sufficient to state that the two specimens of A. schmidti show no meristic 
differences from A. catbo nor any measurable differences in body proportions. The 
shorter dorsal rays noted by Saemundsson are merely broken ; the intangible 
differences in the contour of the head are due to variations of desiccation and fixation, 
and may be observed in some of the fishes on the Funchal Market slabs ; the colour 
described with poetic exactitude by Saemundsson is merely that of a living A . carbo 
and changes to a glossy black as a post-mortem effect. 



THE FAMILY TRICHIURIDAE 85 

A. minor Collett is founded on a wretched half-grown fish which had somehow 
contrived to survive the loss of its tail. I have compared the holotype with a 
Madeiran specimen of equivalent snout- vent length ; there are no differences. 

A. microphthalmus Norman has been checked against a similar-sized specimen 
from Madeira ; there are no significant differences. The distension of the branchio- 
stegal region of the holotype, adequately shown in Norman's figure, gives an exag- 
gerated superficial impression of a deeper head and smaller eye. 

Sim (1898) compares a Scottish fish with Day's description of Lepidopus caudattis 
and comments : 

" Now in the specimen under notice there is not the slightest indication of 
such ventral scales, and what is considered a scale by the authors named takes 
the form of a strong, bayonet-shaped spine situated behind the vent, and is an 
inch long." 

Sim clearly had an Aphanopus carbo, at that time unrecognised in the British fauna 
but since found to be common along the 100 fathom line, where it may sometimes 
be taken even by the hundred by vessels trawling for hake. 

I have a monograph in preparation covering the anatomy and biology of this 
species. 

Genus BENTHODESMUS Goode & Bean 
Benthodesmus Goode & Bean, 1882, Proc. U.S. Nat. Mus. 4 : 379. 

Type species Lepidopus elongatus Clarke. Three species. 

Goode & Bean erected this genus on the occasion of their describing a fish from Newfound- 
land which they believed to belong to Clarke's New Zealand species (the holotype of which 
they had not seen) and attributed characters to Benthodesmus additional or contrary to 
those in Clarke's description. In 1895 (Oceanic Ichthyology : 206) they erected a new species 
B. atlanticus on their Newfoundland specimen, leaving the situation that Benthodesmus 
was based on a species which they had not seen. Since the holotype of L. elongatus has 
been lost I propose to request the International Commission on Zoological Nomenclature 
to recognize B. atlanticus G. & B. as the type-species of Benthodesmus, which would at the 
same time provide a more convenient reference point and a more satisfactory indication 
of Goode & Bean's intentions. It is practically certain that the two nominal species will 
eventually be shown to be identical, but for the present I am retaining them both until 
New Zealand material shall be forthcoming. B. atlanticus is a j unior synonym of Aphanopus 
simonyi Steindachner. 

Synonyms 

Lepidopus (non Gouan, 1770) \ Numerous authors ; for references see under synonymies 
Aphanopus (non Lowe, 1839) J of species. 

It has been suggested to me that Benthodesmus should be split and a new genus 
erected on B. tenuis (Giinther). I am strongly opposed to any such action, being of 
the opinion that B. tenuis is the close ancestor of B. elongatus and that it would be 
improper to obscure this close relation in the way proposed. 

In the event of a new genus being recognized there is some possibility of the name 
Scarcina Rafinesque (1810) being already available, with S. argyrea preceding B. 
tenuis. Scarcina has always been regarded as a junior synonym of Lepidopus Gouan 
(1770) and for reasons outlined on p. 94 I prefer to leave it so for the present. 



86 



THE FAMILY TRICHIURIDAE 





the family trichiuridae 87 

Diagnosis : 

(1) Body very elongate ; head-length 7-0-7-6 in standard length 221-591 mm., 
body-depth 23-8-34-4 in same (B. tenuis) or head-length 6-8-7-8 in S.L. 
910-1225 mm., body-depth 21.7-27-0 (B. simonyi). 

(2) Tail 55% (B. tenuis) or 60% (B. simonyi) of S.L. 

(3) Vertebrae 47-52 + 75~8o = 123-131 (B. tenuis) or 52-53 + 101-103 = 
153-156 (B. simonyi). 

(4) Spinous dorsal base half as long as soft dorsal base. 

(5) Dorsal spines 39-42, dorsal soft rays 80-88 (B. tenuis) or dorsal spines 
45-46, dorsal soft rays 102-108 (B. simonyi). 

(6) The number of soft dorsal rays is very close to that of the caudal vertebrae. 
There are very few intercalated interneural elements, which are usually 
toward the beginning or end of the fin. 

(7) Anal spines i -f I, the former extremely minute and completely concealed 
in the adult. I is a delicate cardiform scute with a median keel projecting 
as a short point between the two rounded posterior lobes. 

(8) Anal spine i articulates a short distance in front of I. Their common basal 
element is a complex representing three or more fused elements. The 
interhaemal spine is a thin keel supported by three slender, tubular, 
cartilage-tipped spines (B. tenuis) or is completely wanting (B. simonyi). 
The horizontally-directed basal occupies the length of three vertebral centra. 

(9) A complete external anal fin of 70-76 split but unbranched rays extending 
from the anal spines nearly to the caudal (B. tenuis) or with the anterior 
rays wanting and the external fin posterior and reduced to about 25 rays 
(B. simonyi). 

(10) Ventral fins inserted immediately before perpendicular through anterior 
end of pectoral base (B. tenuis) or immediately behind perpendicular 
through posterior end of pectoral base (B. simonyi). 

(11) Ventral fin I, 1 (soft ray always present ?) ; a scale-like spine and an 
internal rudimentary soft ray shorter than the scale. 

(12) The principal teeth of the premaxillary and dentary series are without 
obvious barbs and without special enamel thickenings at the tips. When 
barbs are present, usually on the premaxillary fangs, they are barely 
perceptible and confined to the hinder edges. The margins of the teeth are 
smooth in both jaws. 

(13) Palatine teeth present in a linear series, exposed (B. tenuis) or concealed 
under mucosa (B. simonyi). 

(14) Principal teeth on first gill-arch few, teeth becoming progressively reduced 
on subsequent arches. 

(15) Intermuscular bones (pleurals and epipleurals) reduced. 

(16) Pigmentation uniform silver sprinkled black. Melanophores thinly 
distributed, except for denser aggregations along lateral line and along 
median dorsal and ventral lines. Dark spots at bases of dorsal and anal 
rays, preceded by large individual stellate melanophores in juveniles. 
Fins shaded with pastel colours. 

ZOOL. 4 , 3. 7 



88 THE FAMILY TRICHIURIDAE 

Key to Species 

Ventral fins inserted before anterior end of pectoral base. 

Dorsal rays 120-133 ; anal elements i + I + 70-76 with external rays through- 
out ; vertebrae 1 23-1 31 ; lateral line strongly developed (less than 15 times in height 
at pectoral) 

Benthodesmus tenuis (Giinther) E. Equatorial Atlantic ; Gulf of 
Mexico ; Indo-Pacific. 
Ventral fins inserted behind posterior end of pectoral base. 

Dorsal rays 147-155 ; anal elements i + I + 91-99 with external rays substan- 
tially confined to posterior third ; vertebrae 153-158 ; lateral line less strongly 
developed (more than 20 times in height at pectoral) 

Benthodesmus elongatus (Clarke) New Zealand ; Australia; S. E. 

Africa (?) 
Benthodesmus simonyi (Steindachner) N. Atlantic ; N.E. Pacific 

For full discussion and complete bibliographies see : — 

Tucker, 1953, Proc. zool. Soc. Lond. 123 : 1 71-197, pis. and text-figs. 

1955, Bull. Mus. Hist. nat. Belg. 31, No 64 : 1-26, 1 pi. and text figs. 

Benthodesmus elongatus (Clarke) 

Lepidopus caudatus (non Euphrasen, 1788) Hutton, 1872, Fishes of New Zealand : 13. 
Lepidopus elongatus Clarke, 1879, Trans. N.Z. Inst. 11 : 294, pi. 14. 

Holotype should be in the Dominion Museum, Wellington, N.Z., but cannot be found 

{fide Mr. J. Moreland in litt.). Type locality Hokitika Beach, W. coast of South Island, 

New Zealand. 
Benthodesmus elongatus (part), Goode & Bean, 1882, Proc. U.S. Nat. Mus. 4 : 380. 
Lepidopus {Benthodesmus) elongatus McCulloch, 1915, Biol Res. " Endeavour," 3 : 152. 
? Benthodesmus atlanticus {non Goode & Bean, 1895) Gilchrist & Von Bonde, 1924, Rep. Fish. 

Mar. biol. Surv. S. Afr. 3, Spec. Rep. 7 : 16. 
? Benthodesmus tenuis {non Giinther, 1877) J. L. B. Smith, 1949, Sea Fishes S. Africa : 312. 

Benthodesmus simonyi (Steindachner) 

(Text-fig. 8). 

? Lepidopus elongatus Clarke, 1879, Trans. N.Z. Inst. 11 : 294, pi. 14. (See above.) 
Benthodesmus elongatus (part), Goode & Bean, 1882, Proc. U.S. Nat. Mus. 4 : 381. 
Aphanopus simonyi Steindachner, 1891, S.B. Akad. Wiss. Wien 100 : 356. 

Holotype should be in the Naturhistorisches Museum, Vienna, but cannot be found 
{fide Dr. D. Kahsbauer, in litt.). Type locality N.E. from S. Cruz de Teneriffe, Canary Is. 
Benthodesmus atlanticus (part) Goode & Bean, 1895, Oceanic Ichthyology : 206. 

Holotype U.S. Nat. Mus. Washington No. 291 16. Type locality W. edge Grand Bank 
of Newfoundland. (The two smaller specimens mentioned are Diplospinus multistriatus 
Maul, fide Dr. Carl L. Hubbs in litt.) 
Lepidopus sp. Vieira, 1895, Ann. Sci. nat. Porto 1 : 165, upper figs. pi. 9 and 10. 
Lepidopus atlanticus, Boulenger, 1899, Ann. Mag. nat. Hist. (7) 3 : 180. 
Lepidopus {Benthodesmus) atlanticus Saemudsson, 1921, Skyrsla um hide islenzka ndtturufraedisf- 

jelag 1919-20 : 37. 
Benthodesmus tenuis {non Giinther, 1877) (part) J. L. B. Smith, 1949, Sea Fishes S. Africa : 312. 

(Figure copy of B. atlanticus from G. & B. 1895.) 
Benthodesmus simonyi Maul, 1953, Proc. zool. Soc. Lond. 123 : 167. 



THE FAMILY TRICHI URID AE 89 

Benthodesmus tenuis (Gunther) 
(Text-fig. 9) 

Lepidopus tenuis Gunther, 1877, Ann. Mag. nat. Hist. (4) 20 : 437. 

Lepidopus tenuis Gunther, 1887, " Challenger " Reps. Zool. 22 : 37, pi. 7, fig. B. 

Holotype B.M. (N.H.) No. 1879.5. 14.297. Type locality "Challenger" St. 232, 
35 n' o" N., 139 28' o" E., off Inosima, Sagami Bay, Japan. 
Benthodesmus tenuis, Goode & Bean, 1895, Oceanic Ichthyology : 206. 
Benthodesmus elongatus (non Clarke, 1879) idem. loc. cit. (figure only, a reversed tracing from 

Gunther, 1887). 
Lepidopus aomori Jordan & Snyder, 1901, /. Coll. Sci. Tokyo, 15 : 303. 

Holotype in the Aomori Museum, Japan. Type locality Aomori Bay. 
Benthodesmus benjamini (part) Fowler, 1938, Proc. U.S. Nat. Mus. 85 : 45, fig. 16. 

Holotype U.S. Nat. Mus. No. 98821. Paratypes 98822-5. Type locality " Albatross " 
St. D.5445, off Philippine Is. (The paratype material is contaminated with Diplospinus 
multistriatus Maul, fide Dr. Carl L. Hubbs, in litt.) 
Benthodesmus atlanticus {non Goode & Bean, 1895) Longley & Hildebrand, 1941, Cat. Fish. 

Tortugas : 73. 
? Lepidopus argenteus (non Bonnaterre, 1788) Brauer, 1906, Wiss Ergebd. " Valdivia," 15 : 292, 
taf. 12, fig. 3. (Fig. erroneously captioned L. gracilis.) 

Benthodesmus sp. incertae sedis 

Lepidopus tenuis (? non Gunther, 1877) Franz, 1910, Abh. Bayer. Akad. 4 Suppl. Bd. 1 : 56. 
(Locality Uraga Channel, Japan.) 

On the information available this specimen cannot be assigned with certainty to 
either B. simonyi or B. tenuis. I do not believe it to be a new species, nor do I accept 
Franz's opinion that it justifies regarding this genus as containing one world-wide 
species. 

Subfamily LEPIDOPODINAE Gill 

Lepidopodinae Gill, 1863, Proc. Acad. nat. Sci. Philad. 1863 : 227. 
Type genus Lepidopus Gouan. 

Genera now recognised. — Lepidopus Gouan ; Evoxymetopon (Poey) Gill ; 
Eupleurogrammus Gill ; Assurger Whitley ; Tentoriceps Whitley. 

Diagnosis : 

(Note. — Since there is considerable diversity among the genera of Lepidopodinae 
and since, through inadequate descriptions and lack of study-material, certain 
characters have not been verified in Evoxymetopon, Assurger and Tentoriceps, it is 
necessary to introduce qualifications into the following diagnosis. For this purpose 
the abbreviations Lep., Evox., Eupl. Ass., & Tent, have been used for the generic 
names). 

A. Slope of snout variable, gentle to steep ; orbits barely entering upper 
profile of head (Lep.) or more or less remote from it (all other genera) ; 
posterior confluence of frontal ridges elevated to support a sagittal crest at 
the nape which (in all genera except Lep.) is continued forward along the 
snout as a ridge-like elevation of the entire ethmofrontal region. 



9 o THE FAMILY TRICHIURIDAE 

B. Cartilaginous protuberance at mandibular symphysis weak or absent ; a 
small, soft projection at the tip of the snout. 

C. Lower hind margin of operculum markedly convex. 

D. Teeth of main series without barbs. (Lep., EupL, Evox., Tent. Ass.). 
(Fangs slightly barbed in Lep.). 

E. Teeth on palatines in a linear series. (Lep., Evox., Eupl.) 

F. Lateral line descending gently from the shoulder and running in a median 
or sub-median postiion along the body, i.e. distance between lateral line 
and ventral profile at anus much more than half distance between lateral 
line and dorsal. 

G. Spinous dorsal fin short, with 10 (Evox.), 9 (Lep.) or 3 (Eupl.) rays. Spinous 
and soft dorsals continuous, without any intervening notch. 

H. Soft dorsal rays precisely corresponding to adjacent caudal vertebrae, each 
basal and interneural element being related to a neural spine. (Lep., Eupl.) 

I. Spinous anal i (Lep., Eupl.) -f I (all genera) ; anterior soft anal rays not 
penetrating skin (Lep. Eupl. Evox. Ass.) and external and functional fin 
effectively confined to posterior ca. 20 rays, or (in Eupl.) absent. 

J. Terminations of dorsal and anal fins sub-opposite (Lep., Evox. Ass.) or anal 
extending alightly beyond dorsal (Eupl). 

K. Caudal fin present, small, normal, forked (Lep., Evox., Ass.) or absent (Eupl., 
Tent.) 

L. Ventral fins always present, composed each of a scale-like spine and some- 
times at least (Lep.) an internal rudimentary soft ray ; insertion retarded, 
1 to 5 eye-diameters behind posterior end of pectoral base. 

M. Pyloric caeca ca. 24 (Lep., Eupl.) 

Osteological literature 

Giinther, i860, Cat. Fish. B.M. 2 : 345-346 (short desc. Lepidopus). 

Starks, 191 1, Stanford Univ. Publ. 5 : 17-26, pi. (skull of Lepidopus). 

Tucker, 1953, Proc. zool. Soc. Lond. 123 : 196, pis. (paired fins and anal of Lepidopus). 

Literature on young stages 

Delsman, 1927, Treubia 9, Livr. 4 : 338 (Eupleurogrammus eggs and larvae). 
Regan, 1916, Sci. Rep. Brit. Antarct. Exped. Zool. 1 : 144, pi. 8 (young Lepidopus). 
Strubberg, 1918, Rep. Dan. Oceanogr. Exped. 2 Biol . A . 6 . II : 7-16 (life-history of Lepidopus) . 

Genus LEPIDOPUS Gouan 

Lepidopus Gouan, 1770, Hist. Piscium : 107, 185, Tab. 1, fig. 4. 

No type species designated. Two species. 

The earliest available binomen is L. argenteus Bonnaterre, 1788, Encycl. Meth. Zool. 
Icht. : 58, pi. 87, fig. 364. Bonnaterre's figure is an accurate reversed tracing of Gouan's 
caricature, but L. argenteus is a synonym, and almost certainly a junior synonym, of 
Trichiurus caudatus Euphrasen, 1788, Handl. K. Vetensk. Akad. Stockholm 9 : 52, tab. 9, 
fig. 2. 

Euphrasen's paper appears in the section of the Handl. K. Vetensk. Akad. for Jan., Feb., 
Mar., 1788, the sections having been issued quarterly with separate title-pages though 
paginated in annual volumes. 



THE FAMILY TRICHIURIDAE 



9r 




92 THE FAMILY TRICHIURIDAE 

In their third and final attempt to establish the dates of publication of the parts of the 
Encyclopedie Methodique, Sherborn & Woodward, 1906, Ann. Mag. nat. Hist. (7) 17 : 578 
could establish nothing more precise concerning Bonnaterre's Ichtyologie than that it 
appeared in livraison 28 of the Encyclopedie issued sometime in 1788. Since, however, 
the livraisons were issued in order and the date April, 1788, can be assigned to livraison 26 
the balance of probability favours Euphrasen's publication as the earlier one. Following 
the nomenclatorial orgy at the earlier part of the nineteenth century Euphrasen's name 
has been the more generally used. 

Synonyms 

(The full references to the following are given in the synonymy of Lepidopus caudatus, p. 93). 
Trichiurus (non Linnaeus, 1758) Vandelli, 1797 ; Holten, 1802. 

Vandellius Shaw, 1803. Type species Vandellius lusitanicus Shaw [ex Vandelli MS.). 
Ziphotheca Montagu, 1809. Type species Ziphotheca tetradens Montagu. 

X ft fh > variant spellings by later authors. 

(non Zyphotheca Swainson, 1839.) 

? Scarcina Rafinesque, 18 10. Type species Scarcina argyrea Rafinesque. 

Diagnosis (based on L. caudatus) : 

(1) Body elongate, head 5-8-7-1 in standard length, greatest depth 10-8-18-3 
in standard length (57-1224 mm.) 

(2) Upper profile of head oblique-concave, rising at about 25 to the longitudinal 
axis from above the snout tip to behind the orbits and thereafter more 
steeply to the dorsal origin ; straight before the orbits. Ethmo-frontal 
region not elevated, posterior confluence of frontal crests strongly elevated. 
Interorbital slightly concave with very low longitudinal ridges. 

(3) Orbit large, 4-9-5-6 in head, touching dorsal profile. 

(4) Dorsal IX, 90-96 ; aggregate 99-105. The first dorsal spine is not enlarged, 
save as a transient larval character. 

(5) Anal spines i + I ; I is a small triangular scale 2 or more in the pupil. 

(6) Anal fin elements i + I -f- 61-64 I anterior rays reduced or absent, 
posterior 20-24 rays supporting fin. 

(7) Posterior end of operculum a broadly rounded point, barely reaching to 
anterior end of pectoral base. 

(8) Ventral fins present, scale-like, inserted an eye-diameter behind the 
posterior end of the pectoral base. 

(9) Caudal fin present. 

(10) Vertebrae 41 + 70-73 = 111-113. 

Key to Species 
Dorsal rays 99-105 ; external anal fin reaching only half way to vent. D.IX, 90-96 ; 
anal elements i + I + 61-64 (the last 20-24 only being external fin-supporting rays) » 
vertebrae 41 + 70-73. 

Head 5 -8-7- 1 in standard length 57-1224 mm. ; depth 14-4 (-i8-3)-io-8 in same ; 
eye 4-9-5*6 in head. 

Ventral fins I-i (1 is an internal rudiment only 1 mm. long in the adult fish), inserted 
an eye-diameter behind the pectoral base ; anal spine I is a small triangular scale, 
less that the pupil. Pyloric caeca 20 +. Colour uniform silvery. 

Lepidopus caudatus (Euphrasen). 
Atlantic, Mediterranean, S. Indian Ocean, S, Pacific, 



THE FAMILY TRICHIURIDAE 



93 



Dorsal rays 82-87 I external anal fin reaching to vent. 

Analysis of dorsal spines and rays not known ; external anal i + I + 45-5$ ; 
vertebrae unknown. 

For body proportions see discussion. 

Ventral fins I-i inserted on or immediately behind perpendicular through posterior 
end of pectoral base ; anal spine I is long, keeled, about three-quarters the diameter 
of the eye. Pyloric caeca unknown. 

This compromise description, based on Jordan & Evermann (1898) and Brauer 
(1906), may include two species or one, of uncertain systematic position, and without 
valid name(s). The discussion on pp. 95-7 explains this unhappy situation. 

" Lepidopus Xantusi " Goode & Bean 
California, Gulf of Guinea. 

Lepidopus caudatus (Euphrasen) 
(Text-fig. 10). 

Trichiurus caudatus Euphrasen, 1788, Handl. K. Vetensk. Akad. Stockholm, 9 : 52, tab. 9, fig. 2. 

Holotype in Alstromerika Museum ? Type locality Cape of Good Hope. 
Lepidopus caudatus, White, 1851, List Brit. Anim. B.M. 8 Fish : 32. 
Lepidopus argenteus Bonnaterre, 1788, Encycl. Meth. Zool. Ichth. : 58, pi. 87, fig. 364. 

(Ex Gouan, 1770.) (See note under Lepidopus p. 90.) There is a partial confusion 
with Lepturus argenteus Linnaeus, 1754 (= Trichiurus) in the text. 
Trichiurus ensiformis Vandelli, 1797, Mem. Acad. Sci. Lisboa, 1 : 70 (nomen nudum). 

(id. fide Nobre, 1935, Faun. Mar. Portugal, 1 Vert. : 260). 
Lepidopus ensiformis, Swainson, 1839, Lard. Cab. Cycl. Fish. 2 : 254. 
Lepidopus gouanianus Lac6pede, 1800, Hist. nat. Poissons 2 : 519. 

(Ex Gouan, 1770.) 
Lepidopus gouani Bloch & Schneider, 1801, Syst. Ichth. 1 : 239, tab. 53, lower fig. 

(Ex Gouan, 1770.) 
Trichiurus gladius Holten, 1802, Skr. nat.-Selsk. Kbh. 5, Heft 2 : 23, Tab. 2, fig. 1. 

Holotype in Copenhagen Museum ? Type locality Portugal. (I am doubtful whether 
this name should not perhaps be attributed to Abildgaard.) 
Vandellius lusitanicus Shaw, 1803, Gen. Zool. Pise. 4 (2) : 199. 

(Ex Vandelli MS.) 
Lepidopus lusitanicus, Leach, 18 15, Zool. Misc. 2 : 7, pi. 62. 
Ziphotheca tetradens Montagu, 1809, Mem. Werner. N. H. Soc. 1 : 81. 

Holotype B.M. (N.H.) No. 1955.6.2. 1. Type locality English Coast. 
Lepidopus tetradens, Fleming, 1828, Hist. Brit. Anim. : 205. 
Lepidopus peronii Risso, 1810, Ichth. Nice : 148, PI. 5, fig. 18. 

Type locality Nice. 
? Scarcina argyrea Rafinesque, 1810, Car. n. gen. : 20, pi. 7, fig. 1. 

Type locality Sicily. 
? Lepidopus argyreus, Cuvier, 1829, Regne Animal 2 Ed. 2 : 217. 
Lepidopus gov anianus Risso, 1826, Hist. Nat. 3 : 290. 

(Ex Gouan, 1770.) 
Lepidopus lex Phillips, 1932, N.Z. Journ. Tech. 13 : 232. 

Syntypes in Dominion Museum, Wellington ? Type locality New Zealand. (Lepidopus 
caudatus of other New Zealand authors ; non L. caudatus Hutton, 1872, Fishes N.Z. : 13, 
who had Benthodesmus elongatus (Clarke).) 
Aphanopus cafbo (non Lowe, 1839) (part) Norman, 1937, in Fraser and Norman, Giant Fishes, 

Whales and Dolphins : 140. 
non Lepidopus caudatus Sim, 1898, Ann. Scot. nat. Hist. 1898 : 53 (mis-identification of Aphanopus 

carbo Lowe). 
non Lepidopus elongatus Clarke (1879) ; McCulloch (191 5) (see Benthodesmus elongatus). 



94 



THE FAMILY TRICHIURIDAE 



non Lepidopus sp. Vieira (1895). \ (See B simonyi \ 

non Lepidopus atlanticus, Boulenger (1899) ; Saemundsson (1921) J v ' ' 

non Lepidopus aomori Jordan & Snyder (1901) 1 ,„ _ ., , . . 

r I.* . . ^ t *\ > (See Benthodesmus tenuis.) 

non Lepidopus argenteus Brauer (1906) J v ' 

The nineteenth century synonyms listed above have been pretty generally accepted; 
I have verified each of them, so far as the accompanying data allow, and do not propose 
to attempt any individual justifications in the present short summary. Only Scarcina 
argyrea Rafinesque (1810) calls for any urgent comment. This name has been copied 
as a synonym of Lepidopus caudatus by many authors, but the figure shows a head 
and body-form very reminiscent of a Benthodesmus and the stated dorsal count (125) 
falls within the range of B. tenuis (Giinther) and is well above the D. 99-1 05 found in 
L. caudatus. The anal count of 15 and the anal fin as figured are, however, quite 
like Lepidopus. Since Benthodesmus is not yet known from the Mediterranean it is 
better to regard Rafinesque's as an inaccurate impression of L. caudatus for the 
present. Should B. tenuis be found in the Mediterranean Scarcina argyrea will have 
to be considered as a senior synonym and it may be thought desirable to invoke the 
Plenary Powers of the International Commission in order to suppress it. Scarcina 
would also precede Benthodesmus. 

Norman (1937) mentions Aphanopus carbo as being " not uncommon in the fish 
markets of the Mediterranean ". In an intensive study of A . carbo I have so far found 
nothing to confirm this statement, which may have been made through some confusion 
of vernacular names. Thus the Portuguese and Madeiran fishermen call A. carbo 
" O Peixe Espada preta " and L. caudatus " O Peixe Espada branca " (Black and 
White Scabbard-fishes, respectively), and in both cases " Peixe-espada " or " Espada" 
for short. 

Phillipps (1932) : 

" examined several frost-fish and found consistent, though slight, differences 
between the New Zealand and Atlantic species . . . 3 to 4 less rays in the dorsal 
fin, 3 or 4 less anal rays, and a total length of head under 7 in total length. 
Goode and Bean's figure shows a species with a longer head, and no dorsal 
spines of greater length than the diameter of the eye. In the New Zealand fish 
the height of the sixth dorsal ray is 5 in the head while in the European fish 
the height of this ray is about 8 or more in the head. The tail of the New Zealand 
frost-fish is not so deeply emarginate and agrees more nearly with that of 
Evoxymetopon taeniatus figured by Goode and Bean." 

Phillipps is presumably referring to Goode & Bean (1895) Oceanic Ichthyology, 
Plate 58, figs. 213 and 214. I have dealt in some detail with the identification of 
fig. 213 under " Lepidopus Xantusi " (p. 96 q.v.) and so for the present it is 
sufficient to state that this figure is a poorish figure of an apparent young Lepidopus 
caudatus and not a very satisfactory basis for any comparison. The head in Goode 
& Bean's figure goes about 7-5 in the total length and is therefore shorter, not longer 
as stated by Phillipps, and typical of a juvenile as opposed both to post-larval and 
adult specimens. The dorsal spines and tail of Goode & Bean's figure are useless as 
evidence. 



THE FAMILY TRICHIURIDAE 



95 



Comparing specimens of as nearly equivalent size as possible I obtain the following 
results : 

Table II. 

Lepidopus caudatus. 



No 



Standard length . 
Head in S.L. 
Depth in S.L. 
Eye in head length 
6th dorsal spine in H.L 
Dorsal count 
Anal count . 



Lisbon. 

1860.4.22.69. 

1 142 mm. 

7-13 

i5'43 

4-92 

5-33 
D.IX, 96 

i+I +40+24 



New Zealand. 

No. 1903.4.30.29. 

1224 mm. 

6-65 

10-83 

5-4i 

5-25 

D.IX, 90 

A. i+I +41 +20 



Apart from the greater depth of the body, in part attributable to age, the New 
Zealand specimen appears to show only the trivial differences to be expected in 
material of a widely ranging pelagic fish taken from the extreme limits of its distribu- 
tion. The variations are no greater than those found in Trichiurid species, of which I 
have been able to study substantial samples and accordingly I am not at present 
prepared to accept Lepidopus lex Phillipps as distinct from L. caudatus (Euphrasen). 
If, however, the separation of L. lex should be considered justified an interesting 
situation arises. Since the type locality of L. caudatus is the Cape of Good Hope it is 
likely that the antipodal forms will be conspecific but distinct from those of the E. 
Atlantic and Mediterranean. L. lex would therefore still fall as a synonym of L. 
caudatus, but L. argenteus Bonnaterre (1788) would have to be revived. A further 
complication would arise in that the Lepidopus caudatus figured by Goode & Bean 
(1895) and uncertainly associated with the holotype of L. xantusi G. & B. appears to 
have the lower dorsal count of L. lex also. (Further discussion under L. xantusi, 
p. 96). 

"Lepidopus xantusi" Goode & Bean 

Lepidopus caudatus (? non Euphrasen) Jordan & Gilbert, 1882, Proc. U.S. Nat. Mus. 5 : 358. 
Lepidopus caudatus (? non Euphrasen) (part) Goode & Bean, 1895, Oceanic Ichth. : 203, (?) 

fig. 213. 
Lepidopus caudatus (? non Euphrasen) Jordan & Evermann, 1896, Bull. U.S. Nat. Mus. No. 

47 : 886. 
Lepidopus caudatus (? non Euphrasen) Jordan & Evermann, 1900, Bull. U.S. Nat. Mus. No. 

47, (?) pi. 136, fig. 373. 
Lepidopus xantusi Goode & Bean, 1895, Oceanic Ichth. : 519. 

Holotype U.S. Nat. Mus. No. 10115. Type locality Cape San Lucas, California. 
Lepidopus xantusi Jordan & Evermann, 1898, Bull. U.S. Nat. Mus. No. 47 : 2843. 
Lepidopus xantusi Jordan & McGregor, 1899, Rep. U.S. Fish. Comm. 24 (1898) : 276. 
Lepidopus xantusi ? Brauer, 1906, Wiss. Ergebd. " Valdivia," 15 : 291, taf. 12, fig. 2. 

The circumstances surrounding the publication of this species are so wretchedly 
unsatisfactory that a new name will have to be found for it by the first worker able 
to re-describe it from material. 

Jordan & Gilbert (1882) list U.S. Nat. Mus. No. 10115, " One specimen, 10 inches 
long, in poor condition " as " Lepidopus caudatus (Euphr.) White " in a catalogue of 



96 THE FAMILY TRICHIURIDAE 

the fishes collected by one John Xantus at Cape San Lucas, California. (The reader 
should beware confusion with the Joanne Xantus whose Asian collections were pub- 
lished by Karoli.) 

Goode & Bean (1895 : 203) give a description of L. caudatus evidently taken from 
Giinther whose name is, in fact, cited. They then refer to a Xantus specimen and on 
p. 13 of the accompanying Atlas of plates they state that their figure of L. caudatus 
is drawn from U.S. Nat. Mus. No. 10115, collected by John Xantus, off Cape 
St. Lucas. On p. 519 of an appendix to the main text, however, this specimen 
becomes the type of a new species with the brief remark : 

" The specific identity of the fish found at St. Lucas by Xantus is so doubtful 
that we prefer to refer to it as L. Xantusi, new specific name." 

We are left to consider whether Article 21 of the International Rules has been complied 
with ; on the text alone L. Xantusi is a nomen nudum and may be saved only by the 
figure, to be discussed presently. 

Jordan & Evermann (1896) give the Giinther-Goode & Bean version of L. caudatus 
(with an addition of pure Giinther) and conclude by assigning the Xantus specimen 
once again to L. caudatus. Jordan & Evermann (1898) have realized that L. Xantusi 
exists and that somebody should give a description of it, but instead of describing 
it from the holotype (10 inches S.L.) they elect to do so from a second Cape San 
Lucas specimen which is more portable (5 J inches S.L.). Jordan & Evermann (1900), 
however, continue to publish Goode & Bean's original figure of the supposed holotype 
of L. Xantusi still with the legend " L. caudatus ". 

The figure published by Goode & Bean has no scale of magnification nor do these 
authors anywhere state the size of their specimen ; for that we have to return to 
Jordan & Gilbert (1882). Moreover the drawing has the tail nicely curved, an 
effective obstruction to accurate measurement of standard length. I derive the 
following data : 

Radial Formula D.99 ; A. (external) 18. 

(mm.) 
Measured distance from snout tip to D. 30 . . . . .125 

Estimated distance from D.30 to D.70 (taken as 4 x mean distance) 
D. 20-30 and D. 70-80) ........ 120 

Measured distance from D.70 to tip caudal peduncle ... 79 

Whence Estimated standard length of figure ...... 324 

Head in S.L. 7-3 ; depth in S.L. 18 • 6 ; eye in head 5-7 ; snout in head 3*1 
Insertion of ventral fins an eye-diameter behind pectoral base. 

But these are the counts found in Lepidopus caudatus (Euphrasen) and these the 
body-proportions of a young fish of that species ! We are therefore driven to one of 
two conclusions : 

Either (1) The figure is drawn, by some accident, from a specimen other than 
the holotype of L. Xantusi Goode & Bean. In this case the name L. xantusi 
Goode & Bean falls as a synonym of L. caudatus (Euphrasen) ; whatever the 



THE FAMILY TRICHIURIDAE 97 

identity of the Xantus specimen, no "definition or description " have been pub- 
lished, nor any figure of that specimen. Further, although Jordan & Evermann 
(1898) and Brauer (1906) give adequate characterisations of a species distinct from 
L. caudatus (Euphrasen) under the name L. Xantusi, their name must fall as a 
homonym of L. Xantusi Goode & Bean under Article 35 of the Rules. 

or (2) The figure is drawn from U.S. Nat. Mus. No. 10115 as stated and represents 
the holotype of L. Xantusi G. & B. In this case L. Xantusi again falls as a synonym 
of L. caudatus (Euphrasen) and L. Xantusi Jordan & Evermann and L. Xantusi Brauer 
again fall, as homonyms, under Article 35 of the Rules. 

The description by Jordan & Evermann (1898) is repeated verbatim by Jordan & 
McGregor (1899). I give the complete text : 

" Head 4 2/3 in body ; depth 3 in head ; eye 51/3; interorbital space 8 
1/3 ; snout 3 ; maxillary 3 1/3. D.82; ATI, 45. Jaws with long, sharp teeth 
in front, followed by single rows of weaker ones, arranged in groups of twos and 
threes. Height of dorsal, near middle of body, 3 in head. Anal preceded by 2 
scutes, the first minute, the second wide, strongly keeled, its length 3/4 the 
diameter of eye. Pectorals of 12 rays, length 2 in head. Each ventral consists 
of a flat keeled spine followed by a minute ray. This species is known from 2 
small mutilated specimens, both found on the beach near San Jose del Cabo, 
Cape San Lucas. The type was taken by John Xantus, about i860, and recorded 
by Jordan & Gilbert as Lepidopus caudatus. The second, of about the same size 
(5 J inches), was taken by Richard C. McGregor, in 1897. From the latter the 
above account was taken. The species differs from Lepidopus caudatus in the 
much shorter dorsal and longer anal. D.103 ; A. 24. (Named for John Xantus 
de Vesey)." 

Additional data, not provided above, are now needed to decide whether this fish 
may remain in Lepidopus when a new name shall be assigned to it ; at present it 
could as well belong to an Aphanopodine genus as to Lepidopus and may even 
represent a new genus connecting Diplospinus and Lepidopus. 

Brauer (1906) gives a description and figure of " L. Xantusi " from the Gulf of 
Guinea and discusses the difficulties of his identification in face of the above descrip- 
tion. The size is not given, but since a scale of magnification is given for some of the 
other figures on the same plate (though not for this) we may assume Xi, hence 151 mm. 
S.L. Brauer gives D.87 ; A.58 ; head 5-5 in S.L.; depth 15 in S.L. ; eye 5J in head. 
It would help if Jordan & Evermann meant " Head 4 2/3 in body (less head) ", 
i.e. 5 2/3 in S.L. , which would also give depth 17 in S.L. instead of 14. The discrepancies 
between the fin-ray counts are obvious. The figure shows a head about intermediate 
in form between Aphanopus and Lepidopus and ventrals inserted barely behind the 
pectorals, not quite so far retarded as in L. caudatus. Clearly we should know more 
about these specimens. 

Genus EVOXYMETOPON (Poey) Gill. 
Evoxymetopon Poey, in Gill, 1863, Proc. Acad. nat. Sci. Philad. 1863 : 227. 

Type species Evoxymetopon taeniatus (Poey) Gill. Monotypic, or two species. 



9 8 



THE FAMILY TRICHIURIDAE 




the family trichiuridae 99 

Diagnosis : 

(i) Body elongate, head 8 in total length, greatest depth 12-13 in total length. 
(1410-1980 mm.) 

(2) Upper profile of head convex, a steep continuous curve from the tip of 
the snout to the origin of the dorsal set at about 45 to the longitudinal 
axis ; slightly convex before the orbits. Structure of cranial crest un- 
known^ but evidently the ethmo-frontal region and the posterior con- 
fluence of the frontal crests are both elevated. Interorbital strongly convex. 

(3) Orbit large, 5-6 in head length, an eye-diameter ± below the dorsal profile. 

(4) Dorsal X, 77 ; aggregate 87. The first dorsal spine may be enlarged, nearly 
as long as the head. 

(5) Anal spines i(?) -f- I ; I is a keeled scale. 

(6) Analysis of anal fin elements unknown ; the anterior rays, if present, 
appear barely to penetrate the skin while the posterior ca. 20 only are 
fin-supporting rays. 

(7) Posterior end of operculum a broadly rounded point falling less than a 
pectoral base short of the pectoral base. 

(8) Ventral fins present, scale-like, inserted an eye-diameter behind the posterior 
end of the pectoral base. 

(9) Caudal fin present. 

(10) Analysis of vertebrae unknown. 

Evoxymetopon taeniatus (Poey) Gill 
(Text-figs. 11 12 13) 

Evoxymetopon taeniatus Poey, in Gill, 1863, Proc. Acad. nat. Sci. Philad. 1863 : 228. 

Holotype U.S. Nat. Mus. No. 5735. Type locality Havana, Cuba. 
Evoxymetopon taeniatus Poey, 1873, Ann. Soc. Esp. Hist. nat. Madrid, 2 : 77, pi. 5. 
Evoxymetopon taeniatus Goode & Bean, 1895, Oceanic Ichthyology : 204, fig. 214. 
? Evoxymetopon poeyi Giinther, 1887, " Challenger " Reps. Zool. 22 : 39, pi. 43. 

Disposal of holotype unknown. Type locality Mauritius. 

(For Evoxymetopon anzac Alexander see under Assurger, p. 106.) 

It is curious that Poey should have waited ten years before publishing his own 
description and first figure of this species. Goode & Bean give a new figure of the 
holotype but their description appears to be derived entirely from Gill and their only 
original contribution is to confuse Gill's percentages with millimetres and so to mislead 
others into believing that the specimen is only one-fourteenth of its true length. 

Evoxymetopon poeyi, described " with great hesitation ... as a second species" 
was based on a dry skin which Giinther received from Mauritius while his " Challenger' 
Report was passing through the press. The ownership and ultimate destination of 
this specimen are unstated and unknown ; there is certainly no evidence that it ever 
became part of the permanent collections of the British Museum (Natural History). 

The salient characters of these two fishes, as compiled from the literature, are given 
in Table III, from which it is apparent that there is a large measure of agreement 
between them and that the differences are readily attributable to age or sex, damage 
or misinterpretation. 



THE FAMILY TRICHIURIDAE 



The elongated first dorsal spine noted in E. poeyi is a striking feature and apparently 
unique among adult Trichiurids ; though it occurs in the young stages of Lepidopus 
it does so only as a transitory condition and one to be regarded, like the disproportion- 
ate ventral fins, as a flotation device, parallelled among many other young Teleosts and 
without phyletic significance. E. taeniatus and E. poeyi may be female and male of 




Fig. 12. — Evoxymetopon taeniatus (Poey) Gill. Head of holotype, 
1,410 mm. T.L. (from Poey). 




10 CM. 

Fig. 13. — Evoxymetopon poeyi Gunther. Head of holotype, 1,980 mm. S.L. 
(re-drawn, after Gunther ; scale added). 



one species (c.f. Anthias) or there may be growth changes between 1410 and 1981 
mm. length, but a quite likely explanation is that the Cuban specimen may be 
damaged. 

The homologies of the parts in other Trichiuridae studied indicate that the post-anal 
structures probably comprise the usual minute spinule (so far overlooked) and a 



THE FAMILY TRICHIURIDAE 



broad scale having a median depression or keel, the pair articulating with a simple 
or compound basal structure. Experience with damaged Benthodesmus material 
provides a ready explanation of how the discrepancies between the accounts of Gill, 
Poey and Gunther may have arisen. 



Table III. 



Total length 

Greatest height/T.L. 
Head length/T.L. 
Orbit/head 
Dorsal rays 



First dorsal spine 



Anal rays . 



Post-anal scute . 



Ventral insertions 

Vent 

Coloration 



Evoxymetopon taeniatus (Poey) 
Gill. 
14 10 mm. (Poey) 

1/12 (Gill) 
1/8 (Gill) 
1/6 (Gill) 

(D.87. " The first ten dorsal 
spines are undivided ; the rest 
split."— Gill.) 

No special mention in either 
Gill or Poey. 



A. 19. " Anal spines numerous 
. . . mostly minute, free, pos- 
teriorly enlarged, connected 
by the membrane and forming 
a fin" (Gill). 

Upwards of 30 small spines 
figured anterior to the fin 
proper (Goode & Bean). 

" Dagger-shaped spine behind 
the anus " (Gill). 

" A corta distancia posterior del 
ano la pequena escama trian- 
gular y movediza senelada por 
Cuvier en el Lepidopo" (Poey) . 

About 1 \ times the head length 
from the tip of the snout (17^ : 
12— Gill). 

" Submedian " (Gill). 



Silvery, with about six narrow 
reddish bands most distinct 
behind, the first on the ridge 
of the back and the fifth along 
the lateral line " (Gill). 



Evoxy metopon poeyi Gunther. 

78 inches (Gunther) 

(ca. 1981 mm.) 

1/13 or less (Gunther) 

1/8 (Gunther) 

1/5 (Gunther) 

D.93 (Gunther) 



. large, compressed, 
sword-shaped . . . not much 
shorter than the head . . . 
loosely articulated with the 
interneural " (Gunther). 

x + 20 (Gunther) 
. . . anal fin, the rays of 
which begin to be free in the 
posterior third of its extent " 
(Gunther). 



(Gill's spine) " is entirely cover- 
ed by skin, and consists of 
coalesced and flattened inter- 
haemai elements ... a single 
oval scale slightly bent along 
the middle occupies the space 
at a short distance behind the 
vent " (Gunther). 

About 1$ times the head length 
from the tip of the snout (313 : 
240 — Giinther's fig.). 

" Somewhat in advance of the 
middle of the total length " 
(Gunther) . 

" Uniform silvery " (Gunther). 



102 THE FAMILY TRICHIURIDAE 

The upper profile of the head in E. poeyi does not rise quite as steeply as in E. 
taeniatus (a condition apparently related to allometric growth of the jaws) and the 
whole head is less plump in appearance. Here, again, one recalls post-mortem changes 
witnessed in freshly caught Aphanopus carbo off Madeira, as well as the fact that 
E. poeyi is figured from a dry skin, and accordingly one discounts the differences. 

Gill (1863) alludes to a Scottish specimen referred by Hoy to Trichiurus lepturus 
(there were actually two) and suggests that it may have been an Evoxymetopon. 
Evoxymetopon has never been taken in British waters and Hoy's specimens must be 
referred probably to Trachypterus or Regalecus. 

Although the osteology of Evoxymetopon is unknown it is certain that the ethmo- 
frontal region of the skull, together with the posterior confluence of the frontal ridges 
must be elevated in much the same way as in Eupleurogrammns, but to a greater 
extent. This character apart Evoxymetopon stands fairly close to Lepidopus and is 
very near to the ancestor of Eupleurogrammus. 



Genus EUPLEUROGRAMMUS Gill 

Eupleurogrammus Gill, 1863, Proc. Acad. nat. Sci. Philad. 1863 : 226. 

Type-species Trichiurus muticus Gray. Two species. 
Trichiurus (part). Many authors, from Linnaeus (1758), whose type material of Trichiurus 

lepturus was contaminated with this genus. (See note under Trichiurus, p. 114). 
Enchelyopus (part) Bleeker, 1872, Ned. Tijdschr. Dierk. 4 (1872) : 131. 

Diagnosis : 

(1) Body very elongate, head 9-4-11-2 in total length, greatest depth (in 
region of vent) 14-7-17-6 in total length (273-617 mm.) 

(2) Upper profile of head oblique to very slightly concave, rising from the tip 
of the snout in a line set at about 30 to the longitudinal axis and quite 
straight before the orbits. Cranial crest formed by elevation of ethmo- 
frontal region and of the posterior confluence of the frontal crests. Inter- 
orbital strongly convex. 

(3) Orbit small, 6-0-7-8 in head, \ to \ an eye-diameter below the dorsal 
profile. 

(4) Dorsal III, 123-131 or III, 143-147 ; aggregate 126-150. First dorsal 
spine not enlarged. 

(5) Anal spines i -f- I ; I is a small triangular scale. 

(6) Anal fin elements i -f I + n 4-1 21 ; the external fin is entirely suppressed 
and the ventral profile smooth. 

(7) Posterior end of operculum a rounded point, overlying middle of pectoral 
fin and base. 

(8) Ventral fins present, scale-like, inserted about 5 eye-diameters behind the 
posterior end of the pectoral base. 

(9) Caudal fin absent. 

(10) Vertebrae 32-35 + 125-128 = 157-162 or 41 + 150-151 = 191-192. 



THE FAMILY TRICHIURIDAE 103 

Key to Species. 
Anal origin below D. 3 3-3 7 

D.III, 123-131 ; Vertebrae 32-35 + 125-128 = 157-162 

Eupleurogrammus intermedins (Gray) 
Indo-Pacific. 
Anal origin below D. 4 1-4 2 

D.III, 143-147 ; Vertebrae 41 + 150-151 = 191-192 

Eupleurogrammus muticus (Gray) 
Indo-Pacific. 

Eupleurogrammus intermedius (Gray) 
(Text-fig. 14) 

Trichiurus intermedins Gray, 1831, Zoo. Misc. 1 : 10. 

Syn-types (3) B.M. (N.H.) No. 1869.3. 19.76. Type locality Chusan. 

Trichiurus muticus (non Gray) numerous authors. (Incorrect deductions from Gray's original 
description or from following Gunther, i860, Cat. Fish. B.M. 2 : 348 ; no new material 
involved.) 

Trichiurus medius Griffith, 1834, Cuvier's Anim. Kingd. Pisces : 349 (nom. emend, from Gray). 

Trichiurus savala {non Cuvier, 1829) (part) Bleeker, 1852, Verh. Bat. Gen. 24 Makr. : 41. (Deter- 
mination altered to T. glossodon by Bleeker, see below.) 

Trichiwrus glossodon Bleeker, i860, Acta. Soc. Indo-Neerl. 8. Dertiende Bijdr. Visch. Borneo : 

38. 

? Syn-types, in Leiden Museum and in British Museum (Natural History), B.M. (N.H.) 

No. 1880. 4. 21. 1 19. Type localities Java, Sumatra, Singapore, Bintang, Borneo. 
Trichiurus glossodon De Beaufort, 1951, Fish. Indo-Austr. Archip. 9 : 190. (Bleeker's material 

re-examined.) 
Trichiurus glossodon Delsman, 1927, Treubia 9, livr. 4 : 338. 

Gunther (i860) regarded Trichiurus intermedius Gray as a synonym of T. muticus 
Gray. This error of judgment not only led almost every subsequent worker astray ; 
it also had the practical result that Gray's types in the British Museum (Natural 
History) were not properly recognized and segregated. There is, however, one jar, 
Reg. No. 1860.19.76, containing three specimens and bearing (among others) a label 
in an old hand stating : 

" Trichiurus intermedius 
Chusan. E. I. Company." 

A second label, written in ink on paint, changes the identification to Trichiurus 
muticus and a third, overlying both, adds the Register number and changes the source 
to " Dr. Cantor's Colin." It is not possible to reconcile this material with any entry 
in Gunther (i860), but there seems no doubt that these are the syntypes of T. 
intermedius Gray, both from their apparent history and their study. 

Accordingly T. intermedins provides one of the major nomenclatorial surprises 
of the present paper. Even as a synonym of T. muticus it would, of course, have 
passed over into Eupleurogrammus, but, as shown in the key and in Table IV. 
T. intermedius proves to be a perfectly valid species. Further, on the evidence of 
a probable syntype of T. glossodon Bleeker and on De Beaufort's (1951) re-description 
of other presumed type-material of T. glossodon at Leiden, it becomes apparent that 
the more widely-recognized T. glossodon is only a synonym of T. intermedius, as 

ZOOL. 4, 3. 8 



io4 



THE FAMILY TRICHIURIDAE 




o 

tO 

H 

to vO 

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• o 

d ■» 

ts ^ 
S CO 



oo 00 

s o 

s? CO 



S CO 

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THE FAMILY 


TRICHIURIDAE 




105 








Table IV. 






A. 

origin 




Standard 


Head 


Depth 


Eye 








Length 


in 


in 


in 


Fin-ray 


counts : 


below 


Vertebral 




(mm.). 


S.L. 


S.L. 


head. 


D. 


A. 


D. 


counts. 


\ Eupleurogrammus intermedins : 


















Trichiurus intermedins Gray 


















Syn-types : Chusan : East f 


273 


10 11 


. 17-61 


. 6 00 . 


HI, 130 


i + 1 + 120 


• 35-6 . 


34 + 128=162 


India Co. Reg. No. B.M. <{ 
(N.H.) 1860.3. 19.76 t 


308 . 


977 


17-11 


. 656 . 


III, 128 


i + I + 116 


• 37 • 


35 + 127=162 


336 


1029 


. 1639 


. 6-56 . 


in; 131 


i + I + 114 


• 35 • 


32 + 127=159 


Trichiurus glossodon Bleeker 


















? Syn-type : No. loc. : Bleeker 


















Coll. Reg. No. B.M. (N.H.) 


















1880. 4. 21 .119 


427 


9-47 


• 15 -oo 


. 6-42 . 


in, 123 


i + I + 114 


• 33 • 


32 + 125 = 157 



ca. 




ca. 










. 10-26 


14-69 


7-0 . 


HI, 143 


i + I + 120 . 


41 


. 41+151 = 192 


. 11-21 


15-42 


7-85 • 


HI, 147 


i + I + I2I . 


42 


. 41+150=191 



Eupleurogrammus muticus : 
Trichiurus muticus Gray 
Holotype : India : Hard- 
wicke, Reg. No. B.M. (N.H.) 
I955-5-I3-2 • • .426 

Eupleurogrammus muticus (Gray) 
No data. Reg. No. B.M. 

(N.H.) 1955.6.4. 1 . . 617 

Bleeker (i860) himself suspected, and Delsman (1927). Delsman goes far towards 
recognizing the different vertebral counts in T. muticus and T. glossodon, but in 
the former case he appears to have had the misfortune to select a specimen with 
a broken tail and gives 40-115 = 155. De Beaufort comprehends the affinities of 
T. muticus and T. glossodon, as he shows in his key, but, apparently not having heard 
of Eupleurogrammus, he retains both species among Trichiurus and gives no Eupleuro- 
grammus combinations in his synonymies. 

De Beaufort's counts on the type material of T. glossodon (D.115-120. A. about 90.) 
seem a little on the low side though, in the nature of the material, not disturbingly so. 

Discussion of the relationships of the two species of Eupleurogrammus and of 
their systematic position is continued under the following species. 

Eupleurogrammus muticus (Gray) 
(Text-fig. 15) 

Trichiurus lepturus (part) Linnaeus, 1758, Syst. Nat. Ed. 10 : 246 {fide Lonnberg et at., 1896, K. 

Svensk. Vet.-Akad. Handl. 22 : 40. See note under Trichiurus, p. 114.) 
Trichiurus muticus Gray, 1831, Zool. Misc. 1 : 10. 

Holotype B.M. (N.H.) No. 1955.5. 13.2. Type locality India. 
Eupleurogrammus muticus, Gill, 1863, Proc. Acad. nat. Sci. Philad. 1863 : 226. 
Enchelyopus muticus, Bleeker, 1872, Ned. Tijdschr. 4 : 131. 
non Trichiurus intermedius Gray, 183 1, Zool. Misc. 1 : 10. 

The taxonomy of Eupleurogrammus muticus requires very little comment, except 
a further emphasis on the fact that T. intermedius Gray is not a synonym of it as 
Gunther has led too many to suppose. I have examined the types of both nominal 
species and give the results in Table IV. The holotype of E. muticus has the gill-covers 
partly damaged, but another and better specimen has been available also. 



106 THE FAMILY TRICHIURIDAE 

It has been discussed whether the distinction holds that E. muticus is " burnished 
silver " and E. intermedins " purely silvery "; Delsman and Day pro, De Beaufort 
con. De Beaufort appears to clinch the matter when he says : 

" My specimens " (of muticus) " do not differ in colour from specimens of 
glossodon ' (= intermedius) * collected in the same locality and preserved in the 
same jar." 
The type of silver coloration is, in fact, quite variable in any one species of Trichiurid, 
depending on age, on fixative and preservative, on the amount of oil in the tissues 
(which can import a golden tinge to the silver) and on the fine or coarse grain of the 
guanine itself. 

Despite the superficial resemblance and absence of a caudal fin it seems surprising 
that these two species should ever have been placed in Trichiurus and still more so 
that Gill (1863) should have been content to recognize Eupleurogrammus without 
removing it to the Lepidopodinae. The typically Lepidopodine palatine teeth and 
median lateral line are fundamentally different from those of the Trichiurinae ; to 
these characters are allied a rounded operculum and the presence of ventral fins. 
Further, though the development of the cranial crests is unlike that of Lepidopus 
as of the Trichiurinae, it is very like that of Evoxymetopon, a Lepidopodine which 
Gill had in his hands and classified as such. The dentition of the main series is finer 
than that of any other genus of the Trichiuridae. 

Authors have regarded the ecaudate genera as " degenerate " simply because of 
their lack of a caudal fin. This is a very hasty and unwise opinion : in fact Eupleuro- 
grammus is one of the most advanced. Not only does it display the culminations of 
a number of progressive trends (see pp. 125-8) ; in appearance and structure it 
has the most elegantly streamlined form. The sides of the head and operculum are 
smoothly curved, with none of that chunkyness which occurs in the more primitive 
genera ; the upper and lower profiles of the body are both gently convex ; the dorsal 
is arched and the always untidy anal entirely suppressed ; the point of greatest depth 
has moved back toward the vent ; and a comparison of a radiograph of the skeleton 
with that of, say Nesiarchus or Diplospinus, is like a comparison between the mecha- 
nism of a high-grade chronometer and of a cheap alarm-clock. 

Genus ASSURGER Whitley 

Assurger Whitley, 1933, Rec. Aust. Mus. 19 : 84. 

Type species Evoxymetopon anzac Alexander. Monotypic, Indo-Pacific. 
Diagnosis : 

(1) Body extremely elongate, head 12 in total length, greatest depth 28 in 
total length (ca. 1415 mm.) 

(2) Upper profile of head oblique, rising continuously from the tip of the snout 
in a straight line set at about 25 ° to the longitudinal axis and quite straight 
before the orbits. Structure of cranial crest unknown, but evidently the 
ethmo-frontal region and the posterior confluence of the frontal crests 
are both elevated. Interorbital strongly convex. 

(3) Orbit small, 8 in head length, i an eye-diameter below the dorsal profile 



THE FAMILY TRICHIURIDAE 107 

(4) Analysis of dorsal spines and soft rays unknown ; aggregate ca. 120. 
First dorsal spine not enlarged. 

(5) Anal spines i(?) + I ; I is a large oval scale. 

(6) Analysis of anal fin elements unknown ; only the posterior 14-15 appear 
to be external fin-supporting rays. 

(7) Posterior end of operculum a rounded rectangle, falling about a pectoral 
base-length short of the pectoral base. 

(8) Ventral fins present, scale-like, inserted about 1 J eye-diameters behind the 
posterior end of pectoral base. 

(9) Caudal fin present. 

(10) Analysis of vertebrae unknown. 

Assurger anzac (Alexander) 
(Text-fig. 16 and PL 10). 

Evoxymetopon anzac Alexander, 1916, /. Roy. Soc. W. Aust. 2 : 104, pi. 7. 

Holotype in the Western Australian Museum, Perth. Type locality North Fremantle, 
Western Australia. 
Evoxymetopon anzac Kamohara, 1952, Sci. Rep. Kochi Univ. No. 3 : 31, fig. 26. 
Assurger alexanderi (" nom. emend., as Anzac is not permissible ") Whitley, 1933 ,Rec. Aust. Mus. 
19 : 84. 

(Whitley's emendation is quite unnecessary since a " Recommendation " at the end of 
Article 14 of the " International Rules of Zoological Nomenclature " expressly states : 
"Latinized Greek words or barbarous words may, however, be used. Examples . . . 
ziczac . . .") 

This species is known from Alexander's original and incomplete description and 
figure, the latter a photograph showing the head and the trunk back to the level of 
about the tenth dorsal ray. Whitley, in a general paper of miscellaneous studies, 
erected Assurger apparently on Alexander's account alone and without examination 
of the specimen which, though remote from Sydney, must surely have been more 
easily accessible to Mr. Whitley than to any worker outside Australia. 

The following are all the data that can be extracted from Alexander : 

" B.7 ; D. circa 120 ; A. 14 -f- ; C. 17 ; P. 12. 

Total length 1415 mm., length of head 120 mm., greatest height 50 mm., 
diameter of orbit 15 mm. 

" Unfortunately the fins are a good deal broken, and it is impossible to count 
the rays of either the dorsal or anal with accuracy, no doubt these breakages 
occurred when it was washed ashore, and if the large spine at the commencement 
of the dorsal found in E. fioeyi was ever present it has disappeared. In other 
respects the example agrees in its structural features with Giinther's description, 
the postanal spine is exposed evidently owing to the abrasion of the skin in that 
region and just behind it there is a large oval scale similar to that described and 
figured by Gunther. There is no trace of the six narrow reddish bands which 
Poey describes in E. taeniatus and if one may judge from Goode and Bean's 
figure, the ridge on the forehead is not nearly so high as in that species, but 
agrees with that of E. foeyi." " . , , a bright silvery colour," 



io8 



THE FAMILY TRICH I U RI D A E 




O <U 

»o,g 



G ^ 

it 

8.1 

.—. in 
t3 - — 
§ rt 

£« 

« y; 

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&0 



THE FAMILY TRICHIURIDAE 109 

Alexander's references are to Poey (1873) and Goode & Bean (1895) (given with 
other Evoxymetopon references on p. 99). Although his discussion mentions Gill 
(1863) an d Giinther (1887) also, there is internal evidence in his paper that he cannot 
have examined them all. Gill clearly states that the type of E. taeniatus is nearly 
five feet long (in which he is followed by Giinther) and Poey gives 1410 mm. total 
length, yet Alexander follows the mistake of Goode & Bean, who copy Gill's percentage 
proportions of total length as millimetres, and gives 100 mm. as the total length. 
When, therefore, Alexander denounces Goode & Bean's " very poor figure " of 
E. taeniatus he is complaining of a figure of a fish which he has not seen and which he 
has not compared with Poey's independent drawing of the same specimen. 

Alexander's figure shows the profile of the head rising in practically a straight line 
from the tip of the snout to behind the eye, the slope (with the mouth open) being 
about 25 to the longitudinal axis of the body. The eye lies half its diameter from the 
dorsal profile. The hinder end of the operculum falls about a pectoral base-length 
short of the pectoral fin. The ventral fins are not mentioned ; the photograph shows a 
nondescript median projection before the level of the end of the operculum, certainly 
irrelevant, and a slight indentation, about an eye-diameter behind the pectoral base, 
which is a likely position for the ventrals but quite inconclusive. 

Dr. L. Glauert, Director of the West Australian Museum, has kindly done what he 
could to amplify Alexander's account. He has provided the original photographic 
print from which Alexander's plate was made and this is reproduced, I hope with 
greater clarity than before, as Plate 10 of the present paper. One point which this 
print does clarify is the fact that there are no barbs on the teeth ; Alexander's 
plate may seem to indicate a barb on one of the premaxillary fangs, but this is an 
artifact due to indistinct reproduction of a piece of rubbish on the tooth in question. 
Dr. Glauert gives the eye-diameter as 16 mm. and the length of the head (measured 
from the snout-tip) as 113 mm., whence the ratio eye /head-length must be 1/7 
instead of 1/8 as given by Alexander. Dr. Glauert is unable to add precision to 
Alexander's account of the dorsal fin : "... a University Undergraduate, interested 
in fishes, made an attempt and counted only 127, whereas all those others who made 
the attempt gave from 135 to 142. The explanation is that the dorsal fin was very 
much damaged when the fish reached the Museum ". 

Kamohara (1952) reports one specimen of 2250 mm. from Kochi Market, Japan, 
and gives a small figure but no description. The illustration shows the fish bent into 
an S which prevents measurements of the body proportions and a count of the dorsal 
fin rays, but the general picture agrees with Alexander's description and figure. One 
new fact emerges : the ventral fins are inserted about i| eye-diameters behind the 
pectoral base. 

Since it is evident that the structure of the ethmoid and frontal region of the head 
must be similar to that in Eupleurogrammus and Evoxymetopon I place this fish 
among the Lepidopodinae, among which it may be considered to parallel Benthodesmus 
among the Aphanopodinae. It is at once separable from Eupleurogrammus and 
Tentoriceps by its possession of a caudal fin and from Lepidopus by the form of the head 
and of the elongate body. It differs from Evoxymetopon in the gentler slope of the 
snout, smaller eye (1/7 : 1/6 of the head), elongate body (height 1/28 : 1/12, head 



THE FAMILY TRICHIURIDAE 



1/12 : 1/8 of total length) and higher dorsal count (ca. 120+ : 87). These compari- 
sons are between holotypes of practically identical size (Assurger anzac 1415 mm., 
Evoxymetopon taeniatus 1410 mm. total length). 



Genus TENTORICEPS Whitley 
Tentoriceps Whitley, 1948, Rec. Aust. Mus. 22 : 94. 

Type species Trichiurus cristatus Klunzinger. Monotypic. 

Diagnosis : 

(1) Body extremely elongate, head 9 in total length, greatest depth 20-24 in 
total length (ca. 418 mm.) 

(2) Upper profile of head convex, a continuous curve rising from the tip of 
the snout at about 30 to the longitudinal axis and markedly convex before 
the orbits. Structure of cranial crest unknown, but evidently the ethmo- 
frontal region and the posterior confluence of the frontal crests are both 
elevated, the former perhaps disproportionately so. Interorbital convex. 

(3) Orbit large, 5 in head length (Kluzinger description) or 6 (Klunzinger 
figure), 2/3 of an eye-diameter below the dorsal profile. 

(4) Analysis of dorsal spines and soft rays unknown ; aggregate ca. 120. 
First dorsal spine not enlarged. 

(5) Analysis of anal fin elements unknown : " mit rudimentaren, kaum sicht- 

(6) baren Stachelchen ". 

(7) Posterior end of operculum acutely elliptical, reaching to middle of, but 
not concealing, pectoral base. 

(8) Ventral fins present, scale-like, but insertion unknown. 

(9) Caudal fin absent. 

(10) Analysis of vertebrae unknown. 

Tentoriceps cristatus (Klunzinger) 
(Text-fig. 17). 

Trichiurus cristatus Klunzinger, 1884, Fische Rothen Meeres 1 : 120, Taf. 13, fig. 5a. 

Syntypes retained in Klunzinger's private collection, Stuttgart ; eventual disposal 
unknown. Type locality Kosseir, Red Sea coast of Egypt. 
Tentoriceps cristatus, Whitley, 1948, Rec. Aust. Mus. 22 : 94. 

All that is known of this species is contained in Klunzinger's original description 
and figure. I give the complete text : 

" Kopfprofil convex, gratartig, scharf : eine hohe blattartige, bogige Crista 
zieht vom Beginn der Rfickenflosse an fiber Stirn und Schnauze ; den vorderen 
Theil der letztern indess nicht mehr scharfend. Das Auge liegt daher weit unter 
der Profillinie. Bauchflossen wie beim vorigen in Form eines Schuppenpaares 
wie bei b." (b. is Trichiurus muticus Gray, type-species of Eupleurogrammus.) 
" Die Seitenlinie senkt sich sehr allmahlig abwarts und lauft etwas fiber dem 
unteren KorperdritteJ . Afterflosse nur mit rudimentaren, kaum sichtbaren 
Stachelchen. Auge gross, 5 in der Kopflange, Schnauze von doppelter Lange 



THE FAMILY TRICHI URID AE in 

des Auges, Kopf massig lang, 2§ mal so lang als der Korper hoch, 9 in der 
Gesammtlange. Korperhohe 20-24 (letzteres bei Aelteren) in der Gesammtlange, 
Korper also sehr gestreckt. Riickenstrahlen 1 1/4 in der Korperhohe, 3 1/2 in 
der Kopflange, also ziemlich nieder. Brustflossen kurz, 7 in der Kopflange, (wenn 
nicht abgebrochen?). Peitsche kurz, nur von 1/2 Kopflange. Vordere Zahne 
einfach ohne Ansatz. D. c. 120 (?). Neue Art vom Rothen Meer." 

" Von dieser neuen und durch die scharfe, blattartige Kopfgrate gut charak- 
terisirten Art (siehe obige Uebersicht) bekam ich 3 Exemplare bei Koseir, 
ebenfalls aus dem inneren Meer. Farbe silbrig, Ruckenflosse hyalin ". 

The lengths of the three specimens are not given. The head, however, figured " in 
natiirlicher Grosse " is 46-5 mm. in length, (measured from the snout tip) and this 
multiplied by 9 gives ca. 418-5 mm. for the total length of the specimen which is 
likely to have been the largest of the three. 




5 CM. 

Fig. 17. — Tentoriceps cristatus (Klunzinger). Head of syntype ca 418 mm. S.L. (re-drawn 
after Klunzinger (1884), scale added). Some confusion is evident in the representation 
of the nostrils. 



Klunzinger's figure shows only the head, pectoral fin and trunk back to the third 
dorsal ray. Even so the evidence available appears adequate to justify the recognition 
of a distinct species and genus, provided it is all accurately related and represented ; 
relatively small divergences from the published account would involve consideration 
of possible Assurger or Eufleurogrammus spp. Klunzinger's careful consideration 
of the whole genus may justify confidence in his present data. 

It is obvious at the outset that T. cristatus has very little in common with the 
Aphanopodinae. It is likewise certain that, despite its ecaudate condition, it differs 
from the Trichiurinae in the general shape of the head, in the presence of ventral fins, 
in the median position of the lateral line and the absence of barbs from the teeth. 
(Elsewhere Klunzinger properly characterisises the barbed teeth and falling lateral 
line in Trichiurus muticus). 



ii2 THE FAMILY TRICHIURIDAE 

Considered now as a possible Lepidopodine species T. cristatus is quite unlike 
Lepidopus. The form of the upper profile of the head is intermediate between 
Evoxymetopon and Assurger, from both of which our species differs in lacking a 
caudal fin. From Klunzinger's figure it seems that the elevation of the ethmo-frontal 
region has proceeded further than that of the posterior confluence of the frontal 
ridges giving an almost teratological appearance which is quite the reverse of the 
condition in the ecaudate Eupleurogrammus. The hind end of the operculum is a 
rounded point with an extension in relation to the pectoral base intermediate between 
that in the Aphanopodinae and Trichiurinae and unlike the other Lepidopodines. 
The number of dorsal rays (ca. 120) is similar to that in Assurger, but the elongation 
of the body, though considerable, is slightly less (depth 20-24 : 28 in length). The 
number of dorsal spines is unknown, likewise the position of the ventral fin-insertions, 
the condition of the post-anal structures and the number of vertebrae ; nevertheless 
I find it possible to accept Tentoriceps cristatus (Klunzinger) as a valid species and 
genus arising from a Lepidopodine offshoot a little before Assurger. 

Whitley (1948) proposes Tentoriceps with no more than a translation of Klunzinger's 
original description of Trichiurus cristatus, without any indication of the supposed 
discriminant characters and with no reference to his own earlier proposal of Assurger. 
He proposes it in a portmanteau paper of " Studies in Ichthyology " having no 
direct concern with the Red Sea fauna, no special interest in the Trichiuridae nor 
any Australian material of that family requiring comment. Tentoriceps is but 
another of Mr. Whitley's foundlings, casually discovered, capriciously re-baptized 
and callously abandoned, in the hope of adoption or decent interment, on the cold 
doorsteps of systematic ichthyology. 

Subfamily TRICHIURINAE Swainson 

Trachiurinae (evident misprint for Trichiurinae) Swainson, 1839, Nat. Hist. Fish. Amphib. 
Rept. 2 : 254. 

Type genus Trichiurus Linnaeus. 

Lepturinae Gill, 1863, Proc. Acad. nat. Sci. Philad. 1863 : 225. 
Type genus Lepturus Artedi (= Trichiurus L.). 

Genera now recognised : Trichiurus Linnaeus ; Leptur acanthus Fowler. 

Diagnosis : 

A. Slope of snout moderate ; orbits barely entering upper profile of head ; 
posterior confluence of frontal ridges elevated as a sagittal crest at the nape. 

B. Cartilaginous protuberance at mandibular symphysis weak ; a small, soft 
projection at the tip of the snout. 

C. Lower hind margin of operculum more or less concave. 

D. Teeth of main series with barbs. 

E. Palatine teeth minute, in a villiform band. 

F. Lateral line descending steeply from the shoulder and running nearer the 
ventral surface of the body, i.e. distance between lateral line and ventral 
profile at anus slightly less than half distance between lateral line and dorsal. 



THE FAMILY TRICHIURIDAE 113 

G. Spinous dorsal fin very short, with 3 or 4 rays. Spinous and soft dorsals 

continuous, without any intervening notch. 
H. Soft dorsal rays precisely corresponding with adjacent caudal vertebrae, 

each basal and interneural element being related to a neural spine. 
I. Spinous anal i -\- I ; soft anal rays reduced to internal rudiments or wanting 

(Trichiurus) or taking the form of minute pungent spines which definitely 

break the ventral profile (Leptur acanthus) . 
J. Anal fin (i.e., basal elements of anal — see I above) extending well beyond 

dorsal. 
K. Caudal fin and hypurals entirely absent. 
L. Ventral fins and girdle entirely absent. 
M. Pyloric caeca 24, perhaps more. 

Osteological liter atufe 

Giinther, i860, Cat. Fish. B.M. 2 : 343-344. (desc. osteology of Trichiurus). 

Starks, 191 1, Stanford Univ. Pubs. No. 5 : 25-26 (desc. general osteology of Trichiurus, comp . 

with Lepidopus) . 

Gregory, 1933, Trans. Amer. Phil. Soc. 23 : 316, fig. 195 (skull of Trichiurus). 

Literature on young stages 

Delsman, 1927, Treubia 9 : 338. 
Liitken, 1880, K. Dansk. Selsk. Skrift. 12 : 409. 
Nair, 1952, Proc. Indian. Acad. Sci. 35B : 225. 
Tang & Wu, 1936, Lingnan Sci. J. 15 : 651. 

Genus TRICHIUR US Linnaeus 

Trichiurus Linnaeus, 1758, Syst. Nat. Ed. 10 : 246. 

Type species Trichiurus Upturns Linnaeus ex Artedi (see note under T. Upturns). Pro- 
bably monotypic. 

Enchelyopus Klein, 1744, Hist. Piscium : 51. 

Enchelyopus Bleeker, 1862, Versl. Akad. Amsterdam 14 : 109. 

Type species Clupea haumela Forskal. (Also spelt Encheliopus by authors. 
Non Enchelyopus Gronovius, 1763). 

Gymnogaster Gronovius, 1754, Mus. Ichth. 1 : 17. 
Type species Anguilla Jamaicensis Sloane. 

Lepturus Artedi, 1738, Desc. Spec. Pise. : in. 
Type species Lepturus argenteus Artedi. 

Lepturus Gill, 1863, Proc. Acad. nat. Sci. Philad. 1863 : 225. 
(Non Lepturus Moehring, 1758 ; Brisson, 1760.) 

? Diepinotus Rafinesque, 1815, Analyse Nat. : 91 (nom. nud.). (Also spelt Dipinotus by authors.) 
? Symphocles Rafinesque, 18 15, Analyse Nat. : 91 (nom. nud.). 
? Nemochirus Rafinesque, 18 15, Analyse Nat. : 91 (nom. nud.). 

Diagnosis : 

(1) Body-proportions highly variable : Head 7-0-9-4 in length, depth 14-4-21-0 
in length. 

(2) Eye relatively large, 5-0-7-0 in head. 



ii 4 THE FAMILY TRICHIURIDAE 

(3) Dorsal spines III ; D.III, 137 in three specimens of three nominal species 
radiographed. (Published aggregate ranges D. 120-140). 

(4) A. i + I + 105-108. 

(5) Post-anal scute (= anal spine I) not enlarged ; a small, triangular scale, 
less than the pupil. 

(6) First basal element of anal fin slightly enlarged, presumably a compound 
of 2, its interhaemal spine lengthened and slightly thickened. There follows 
a gap of 1 in the series of interhaemal spines, leaving 1 free haemal arch. 

(7) " Soft anal " elements minute spinules which usually do not break the skin 
and which are occasionally absent. The first ca. 60 are directed backwards, 
the last ca. 40 are directed forwards. 

(8) Vertebrae 39-40 + 123-128 = 162-168. 

Probably only one variable species, Trichiurus Upturns L., world-wide except in 
colder regions. 

Trichiurus lepturus Linnaeus 

(Text-fig. 18). 

Trichiurus lepturus (part) Linnaeus (ex Artedi), 1758, Syst. Nat. Ed. 10 : 246. 

Type in the Museum of the Royal University of Upsala. Type locality South Carolina. 

Note. — Lonnberg et al., 1896, K. Svensk. Vet.-Akad. Handl. 22 : 40, state that the 

Linnaean types of " T. lepturus " at Upsala include material of the species now known as 

Eupleurogrammus muticus (Gray). The suggestion that T. lepturus should consequently 

be replaced by T. argenteus Linnaeus, 1754, Mus. Ad. Frid. : 76, pi. 26, fig. 2 is, of course, 

illegal, nor is it really necessary since the situation has never created any practical difficulty. 

Giinther, 1898, Proc. Linn. Soc. Lond. 1898-9 : 29, satisfied himself that the Linnaean 

material in the possession of the Linnaean Society of London is, in fact, T. lepturus, which 

is rendered doubly certain by the fact that it came from Garden's South Carolina collections, 

consignment of 1761. (See also id. ib. : 25.) 

Trichiurus lepturus J. L. B. Smith, 1949, Sea Fish South Africa : 313 ; Okada, 1955, Fishes of 

Japan : 155. 
Trichiurus argenteus Shaw, 1803, Gen. Zool. 4 : 90, pi. 12 (apparently ex Linnaeus, 1754). 
Clupea haumela Forskal, 1775, Descr. Anim. : 72. 

Type not in Herbarium Ichthylogicum Forskalii, Copenhagen (fide N. B. Marshall, 
personal communication) . Type locality Red Sea. 
Trichiurus hamrela Schneider, 1801, Syst. Ichth. : 518 (norn. err.). 
Trichiurus lepturus japonicus Temminck & Schlegel, 1844, Faun. Jap. Pise. : 102, pi. 54. 

Type Leiden Museum No. 2040. Type locality Japan. 
Trichiurus lepturus japonicus Boeseman, 1947, Zool. Meded. 28 : 96 (for Temminck co-author 

p. 2). 
Trichiurus japonicus Bleeker, 1857, Verh. Bat. Gen. 26 : 98. 
Trichiurus japonicus Lin, 1936, Bull. Chekiang Fish. Sta. 2 (5) : 2. 
Trichiurus lajor Bleeker, 1854, Nat. Tijdschr. Ned. Indie 7 : 248. 

Type in Leiden Museum. Type locality Manado, Celebes. (Re-examined by De Beaufort, 
1 95 1, Fish. Indo-Austr. A r chip. 9 : 196.) 
Trichiurus malabaricus Day, 1865, Proc. zool. Soc. Lond. 1865 : 20. 

Holotype B.M. (N.H.) No. 1867.5.30.2. Type locality Madras. (Withdrawn as T. 
haumela by Day, 1876, Fish, India : 201.) 



THE FAMILY TRICHIURIDAE 



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n6 THE FAMILY TRICHIURIDAE 

Trichiurus auriga Klunzinger, 1884, Fische Rothen Meeres 1 : 120, PI. 12, fig. 1. 

Type retained in Klunzinger's private collection, Stuttgart. Now at Stuttgart, Berlin 
or Vienna ? Type locality Kosseir, Red Sea coast of Egypt. 
Trichiurus auriga Weber, 1913, Siboga Fische : 406. 
Trichiurus auriga De Beaufort, 195 1, op. cit. : 196. 
Trichiurus coxii Ramsay & Ogilby, 1887, Proc. Linn. Soc. N.S.W. 1887 2 (2) : 562. 

Holotype Australian Museum, Sydney No. 1. 1342. Type locality Broken Bay, N.S.W. 
? Trichiurus nitens Garman, 1899, Mem. Mus. comp. Zool. Harv. 26 : 69. 

Syntypes (2) in U.S. N.M. ? Type locality coast of Peru. 
? Trichiurus nitens Hubbs & Hubbs, 194 1, Calif. Fish Game 27 : 29. 
? Trichiurus nitens Breder, 1936, Bull. Bingham Ocean. Coll. 2 Art. 3:12. 
non Trichiurus lepturus Mohr, 1786, Forsog til en islandsk Naturh. Kjob. : 63. 
non Trichiurus lepturus Sveinn Palsson, 179 +, J. Naturforsk. Reise Island 1791-97 2. 

(Mis-identifications of Trachypterus sp. Refs. fide Saemundsson, 1926, Fiskarnir : 155. 
Reykjavik). 
non Trichiurus lepturus Hoy, 1815, Trans. Linn. Soc. Lond. 11 : 210-212. 

(Mis-identification of Trachypterus or Regalecus spp.) 
non Trichiurus trimaculatus Giovene, 1829, Mem. Soc. Ital. 20 Pt. 1 : 25. 

(Mis-identification of Trachypterus sp.) 

At the commencement of the present paper Trichiurus seemed likely to give the 
most difficulty ; that promise has been abundantly fulfilled. Trichiurus is a common 
pelagic fish of world-wide distribution, occurring in all but the coldest seas and 
assuming some economic importance in certain areas ; as a consequence it possesses 
a literature as large as that of the rest of the family put together. Much of this work 
is uncritical : species have been recognized on supposed differences of body-proportions 
unrelated to possible ontogenetic changes, geographic variation, or environmental 
effects, or on small differences in fin-ray counts which are difficult to establish 
with any accuracy except in radiographs. Vertebral counts have hardly ever been 
employed. Very often it is found that where a worker has characterised a pair of 
species to his own satisfaction another will reverse the discriminant characters in 
the same pair. 

In dealing with all this intractable material it has seemed useful to take as a 
working hypothesis the theory that we are dealing with one highly variable species. 
If the evolutionary behaviour of the other recent Trichiuridae affords any precedent 
it is one pointing to the evolution of monotypic genera, or of pairs of species having 
sharply discontinuous ranges of meristic counts, not to the subtle distinctions which 
the would-be splitters of Trichiurus postulate. Geographic variation and the increasing 
evidence of environmental effects upon meristic characters must also be taken into 
account. 

The first problem is the identity or distinctness of the Atlantic and Indo-Pacific 
populations; currently recognized as T. lepturus L. and T. haumela (Forskal) 
respectively. The results of the examination of two specimens taken at random 
from the collections of the British Museum (Natural History) are given in Table V. 
They show a precisely coincident dorsal count and anal /vertebral counts differing 
by only 3/2 rays /vertebrae respectively. The differences in body proportions are no 
greater than may be explained by the difference in age. These two specimens show 
that similar Trichiurus occur off Texas and Shanghai almost as well as population 
samples treated with all the apparatus of statistical necromancy. 



THE FAMILY TRICHIURIDAE 



117 





Table 


V 








Trichiurus 




Trichiurus 


Trichiurus 




lepturus. 




" haumela." 


" japonicus." 




Aransas Bay, Texas. 




Shanghai. 


Yenting, Chekiang. 




No. 1948.8.6.795. 


No 


1862. 1 1. 1. 260. 


No. 1925.4.23.5 


Dorsal rays 


D.III, 137 




D.III, 137 


D.III, 137 


Anal rays 


A.i+I+105 




A.i+I+108 


A.i+I+107 


Vertebrae . 


39 + 123 = 162 




46 + 124 = 164 


40+128 = 168 


Standard length . 


545 mm. 




838 mm. 


926 mm. 


Head in S.L. 


7-07 




7-83 


9-17 


Depth in SX. 


16-03 




16-43 


17-47 


Snout-vent in S.L. 


2-79 




3-07 


3-o7 


Head in S-V. 


2-53 




2'55 


2-98 


Depth in S-V. 


5'73 




5*35 


5*68 


Eye in head 


5*92 




6-68 


6-12 


Snout in head 


2-80 




2-77 


2-52 


Mx. in head 


2-48 




2-46 


2*29 



A modern re-assessment of Trichiurus japonicus Temminck & Schlegel, a second 
Pacific form, is given by Boeseman (1947) : 

' The differences between Trichiurus japonicus T. & S. and T. lepturus L. 
as stated by Temminck & Schlegel and Bleeker (I.e.) do not exist in our material. 
A comparison, however, with several specimens of lepturus in our collection (all 
Atlantic) showed a very distinct and constant difference ; the head in all these 
specimens of lepturus is larger, about 7-7*5 in length, while in our specimens of 
japonicus it is 8-1-8-6 in length, consequently considerably smaller. On account 
of this I want to discriminate both species and regard the Japanese specimens 
as the type material of a separate species, Trichiurus japonicus T. & S. Specimen 
no. 2040 I regard as type." 
In opposition to this stands the work of Lin (1936) who comments that : 

" Several authors . . . used to distinguish T. japonicus from T. haumela 
by the shorter head and the smaller eyes, but the series of intermediate forms 
lying between them is so continuous that no clear line can be drawn to separate 
them into two distinct species." 

Lin gives a table of data covering a series of 12 Chinese T. japonicus of 424-1,290 
mm. S.L., with the following ranges : depth in length 14-4-21 ; head in length 7-9-4 ; 
depth in head 1-9-2-3 ; eye in head 5-4-6-4 ; D. 136-140. He notes the difficulties 
which arise through the loss of the tip of the tail and advocates the substitution of 
snout- vent length for the calculation of body-proportions. Oddly enough Lin does 
not take the logical step of substituting the earlier name haumela for japonicus. 

" In reviewing the recorded distribution of T. japonicus and T. haumela, 
it is found that the former species inhabits the Chinese and Japanese coastal 
waters and is neither known to live beyond the Asiatic continental shelf nor 



n8 THE FAMILY TRICHIURIDAE 

southward to the East Indian seas, while the latter is found from the Japanese 
and Chinese seas to the Philippines and Indian Ocean and Archipelago." 

A specimen of T. japonicus (Table V), being duly radiographed, shows a dorsal 
count identical with that of the two specimens of the other two nominal species, an 
intermediate anal count and an increment in caudal vertebrae altogether less than 
the head/body proportion would lead one to expect. I suggest that T. japonicus is, 
in the light of the evidence, no more than an ecotypic form of T. haumela (= lepturus). 

T. lajor Bleeker and T. malabaricus Day have been adequately dealt with by De 
Beaufort (1951) and by Day himself (1876) and as synonyms of T. haumela now need 
no further comment. 

T. auriga Klunzinger, placed very close to T. haumela by Klunzinger (1884) 
himself and founded on a very young specimen (250 mm. S.L.) is probably no more 
than a juvenile of the latter species, though the published illlustration contains 
peculiar features at variance with the description. The only serious difficulty is the 
definitely stated absence of barbs from the teeth. T. auriga has been reported only 
once more, a specimen of 320 mm. from the Timor Sea having been described by 
Weber (1913) and De Beaufort (1951). 

T. coxii Ramsay & Ogilby (1887) contains, on the published account, no differences 
from T. haumela and its authors do not attempt to indicate those characters in which 
they consider it to be divergent. 

Only in the case of T. nitens Garmen (1899) has there been argued any very 
cogent case for specific separation, by Breder (1936) and by Hubbs & Hubbs (1941). 
Breder's conclusions, summarized in the form of a key are: 
A. Dorsal rays never less than 126, usually about 133 ; maxillary 2*2 to 2*5 in head, 

usually about 2*3 (11 specimens, D. 126-137, mean 132*9) . . . lepturus Atlantic 

AA. Dorsal rays never more than 128, usually about 122 ; maxillary 2 "5 to 2 *8 in head, 
usually about 2*6 (36+ specimens, D. 120-128, mean ca. 122*3 

nitens California, Galapagos 

These results must obviously be treated with respect, though not with absolute 
aquiescence. The variation in the maxillary should be stated in relation to the size 
of the fish, for there is some allometry with age. The differences in mean dorsal 
counts are considerable, but if the two samples are drawn from a continuous hypo- 
thetical population of one species, having its origin somewhere in the Indian Ocean 
and extending westwards to the Western Atlantic and eastwards to the Pacific coast 
of America we still have no more than the variation we should expect at the limits 
of that wide range. The satisfactory establishment of T. nitens requires, not compari- 
son with Atlantic material, but demonstration of a non-cline discontinuity across 
the Pacific. 

Having regard therefore to the known variation in the Trichiuridae and to the 
growing literature concerning environmental effects upon fishes, I am more inclined 
to " lump " Trichiurus as one variable species than to " split " and thereby make 
assumptions concerning the genetic distinctness of populations which present 
precedents and evidences of variation do not appear to justify. Such modern authors 
as J. L. B. Smith (1949) and Okada (1955) have apparently arrived at the same 
conclusion, though they do not submit any evidence to support their decisions. 



THE FAMILY TRICHIURIDAE 119 

Genus LEPTURACANTHUS Fowler 

Leptuf 'acanthus (sub-genus of Trichiurus L.) Fowler, 1905, Proc. Acad. nat. Sci. Philad. 1904 : 
770. 

Type species Trichiurus savala Cuvier. Monotypic. 
Trichiurus (part) many earlier authors. 

Diagnosis : 

(1) Body-proportions highly variable : Head 7-4-10-5 in length, depth 14-8- 
19-8 in length. 

(2) Eye relatively small, 6-7-10-0 in head. 

(3) Dorsal spines IV ; D.IV, in in two specimens radiographed, including 
holotype of Trichiurus armatus Gray. (Published aggregate ranges 
D.105-134). 

(4) A.i + I + 72. 

(5) Post-anal scute (== anal spine I) enlarged, as in Aphanopus ; a dagger-like 
spike half the diameter of the eye. 

(6) First basal element of anal fin markedly enlarged, as in Aphanopus, 
presumably a compound of 3, its interhaemal spine likewise lengthened 
and thickened. There follows a gap of 2 in the series of interhaemal spines, 
leaving 2 free haemal arches. 

(7) " Soft anal " elements pungent spinules, definitely breaking the ventral 
profile throughout the length of the fin and all directed backwards. 

(8) Vertebrae 32-35 -f- 124-130 == 159-162. 

One species, Leptur acanthus savala (Cuvier). Indo-Pacific. 

Leptur acanthus savala (Cuvier) 
(Text-fig. 19). 

Trichiurus savala Cuvier, 1829, Regne Animal 2 Ed. 2 : 219. 

Syntypes in Paris Museum, Reg. No. a. 5357-5358. Type locality " Mer des Indes " 
(= Bombay & Malabar). 
Trichiurus [Leptur acanthus) savala Fowler, 1905, Proc. Acad. nat. Sci. Philad. 1904 : 770. 
Trichiurus armatus Gray, 1831, Zool Misc. 1:9; Gray, 1835, Illust. Ind. Zool., pi. 93, fig. 1. 

Holotype B.M. (N.H.) No. 1955.5. 13. 1. Type locality India. 
Trichiurus Roelandti Bleeker, i860, Acta Soc. Indo-Neer. 8 (4) : 30. 

Holotype in Leiden Museum. Type locality Sunda Strait. (Re-examined by De 
Beaufort, 195 1, Fish. Indo-Austr. A r chip. 9 : 194.) 

Trichiurus armatus Gray has for long, and, for once, correctly, been regarded as 
a synonym of this species. De Beaufort has adequately dealt with T. Roelandti 
Bleeker and so the taxonomic situation in this newly-promoted genus Leptur acanthus 
is mercifully straightforward. 

Leptur acanthus is obviously closely related to Trichiurus and as widely separated 
from the other Trichiuridae : any attempt at a natural classification must adequately 
express this situation. In a wide classification Fowler's erection of Leptur acanthus 
as a sub-genus of Trichiurus very adequately did so, but with the exposure of Eupleuro- 
grammus and its removal to the Lepidopodinae the Trichiurinae are left as a very 

ZOOL, 4, 3. 9 



120 THE FAMILY TRICHIURIDAE 

small group. Within this group the divergence between Leptur acanthus savala and 
Trichiurus lepturus is very comparable to that between Aphanopus and Benthodesmus 
and accordingly consistency requires the elevation of Leptur acanthus to full generic 
status. 

THE ORIGIN, EVOLUTION AND CLASSIFICATION 
OF THE TRICHIURIDAE 

Summary of earlier work 

Before summarizing previous opinions on the classification of the Trichiuridae 
it may be useful to indicate the sequence of recognition of the genera now accepted : 

Trichiurus Linnaeus, 1758. 

Lepidopus Gouan, 1770 • 

Aphanopus Lowe, 1839. 

Evoxymetopon (Poey) Gill, 1863. 

Eupleurogrammus Gill, 1863. 

Benthodesmus Goode & Bean, 1882. 

Leptur acanthus Fowler, 1905. 

Assurger Whitley, 1933- 

Diplospinus Maul, 1948. 

Tentoriceps, Whitley, 1948. 

Classification commences, and commences remarkably well, with Cuvier & 
Valenciennes (1831) who recognize the Scombroids as a natural group (Scomberoides) 
containing all the Trichiurids and Gempylids so far known (Lepidopes, Trichiures ; 
Thyrsites, Gempyles) as well as the Tunnies, etc. Their key runs : 

" Tons ou une grande partie des rayons de Vanale reduits a de tres-petites epines. 
Dents des thyrsites et des gempyles. 

Lepidopes. Une petite ecaille au lieu de chaque ventrale ; une 

caudale. 
Trichiures. Point de ventrales ; point de caudale." 

These authors grasp so early the essential relationships of the Trichiuridae and 
Gempylidae : 

" II est impossible de ne pas placer a la suite des gempyles et des thyrsites 
deux genres de poissons qui leur ressemblent presque en toutes choses, si ce 
n'est qu'ils manquent entierement de fausses nageoires et meme de rayons 
mous a leur dorsale ; ce sont les lepidopes et les trichiures, poissons tres- 
remarquables d'ailleurs par leur eclat et par leurs formes singulieres. 

" Leur tete, leurs dents, leur peau, leur squelette, rappellent de tout point les 
genres auxquels nous les associons, et la longueur meme de leur corps en ruban, 
qui les avait fait rapprocher des cepoloides, est deja annoncee par la forme de 
plusieurs gempyles." 

Swainson (1839) was the first to erect a higher taxon for the Trichiurid fishes, 
though he takes a step backwards, making Ammodytes as well as Trichiurus and 
Gempylus members of a sub-family Trichiurinae of the family Coryphaenidae. 



THE FAMILY TRICHIURIDAE 121 

Gunther (i860) attempts no subdivision of his family Trichiuridae, which includes 
not only Aphanopus, Lepidopus and Trichiurus but also the Gempylids Epinnula, 
Dicrotus, Thyrsites and Gempylus. 

Gill (1863) gives a classification recognizably approaching that now advocated, 
though based on inadequate and in part inaccurate premises : 

" I. Dorsal fin undivided. 

A. Tail filiform and finless ....... Lepturinae. 



Lateral line near the abdomen 

Lateral line median .... 

B. Tail with a normally developed and forked fin 
Profile rectilinear and forehead depressed 
Profile high, trenchant and boldly declining 
II. Dorsal fin double ...... 

Teeth of the palate wanting 



Lepturus. 

Eupleurogrammus . 

Lepidopodinae. 

Lepidopus. 

Evoxymetopon. 

Aphanopodinae. 

Aphanopus. 



Johnson (1865) describes a number of Gempylids as Trichiuridae. 
Capello (1868) takes a view of the Trichiuridae equivalent to the Trichiuriformes 
of later authors and recognizes three sub-families : 

C Trichiurus, Eupleurogrammus. 
Trichiurina < Lepidopus, Evoxmyetopon. 

[_ Aphanopus. 
Gempylina . . . . . Gempylus j Prometheus, Epinnula. 

Thyrsitina . . . . . Thyrsites, Dicrotus. 

A division of the Trichiurina similar to Gill's is implicit in the key given. Time, on 
the whole, has dealt more kindly with Capello than did Gunther in the Zoological 
Record. 

Goode & Bean (1895) limit the Trichiuridae to Trichiurus, of which Eupleurogrammus 
is merely " a Chinese form . . . with a single species"! They erect a separate 
family, the Lepidopidae, with two sub-families : 

"I. Dorsal continuous. Teeth on palatines. Ventrals present, scale-like, rudimentary. 

No post- anal spine ......... Lepidopinae. 

(Genera Lepidopus, Evoxymetopon, Benthodesmus.) 
II. Dorsal in two subequal portions, closely contiguous. No teeth on palatines. 

Ventrals absent. A dagger-like post-anal spine .... Aphanopinae. 

(Genus Aphanopus.) 

This pastiche of half-truth and etymological abomination is preceded by one of 
Goode & Bean's self-contradictions (Lepidopidae have " No teeth on palatines "; 
Lepidopinae have " Teeth on palatines "). 

Boulenger (1904) and Goodrich (1909) include both Trichiurids and Gempylids 
in a family Trichiuridae without subdivisions. This grouping becomes the Scombroid 
Division Trichiuriformes of Regan (1909), with two undivided families Trichiuridae 
and Gempylidae in the generally accepted modern sense, as later followed by Jordan 
(1923) and by Berg (1940). 

Starks (191 1), in a classic paper concerned only with the osteology of three genera, 
defines families Gempylidae (Promethichthys) , Lepidopidae (Lepidopus) and Trichi- 

ZOOL. 4, 3. 9 § 



122 THE FAMILY TRICHIURIDAE 

uridae (Trichiurus). He comments that " the descent of the family Trichiuridae 
from the Gempylidae was long ago pointed out " and compares the structure of 
Lepidopus with that of Promethichthys, which latter he rightly regards as too 
specialized to be an ancestral form. He concludes : " This ancestor may have been 
Gempylus, a form which I have been unable to obtain, but showing a development 
towards the elongate forms of Lepidopus and Trichiurus." 

Roule (1927) introduces a little light relief by attempting to place the Iniomous 
Anotopterus among the Trichiuridae. 

Gregory (1933) figures a museum exhibit showing in pictorial form the evolution 
of the Scombroid fishes. Ruvettus, Epinnula, Gempylus and Trichiurus are shown as 
consecutive stages in a linear series, with Lepidopus emerging as a sideshoot between 
Epinnula and Gempylus. In these circumstances the presence of a number of 
apparently undecided fishes swimming in the background to this exhibit occasions no 
surprise. 

Tucker (1953), though not attempting a full classification, draws attention to the 
affinities between Aphanopus and Benthodesmus as contrasted with Lepidopus. 
He shows that Benthodesmus has a differentiated and partly divided dorsal fin like 
that of Aphanopus and demonstrates the significance of the ventral fin-insertions 
and post-anal structures, but, fails to realize that the dorsal fin is differentiated 
throughout the entire family. The error arose through an undue reliance on previous 
literature of the non-Aphanopodinae and a brief study of Lepidopus and Trichiurus 
from radiographs which, for reasons of economy, were fragmentary. Dr. Carl L. 
Hubbs, in litt. kindly drew attention to this mistake. 

Nesiarchus-Diplospinus : the Gempylid-Trichiurid bridge 

Regan (1909) gives the following diagnosis of the Gempylidae which may still 
serve as a basis for comparison with the Trichiuridae (p. 74) : 

" Body oblong or elongate, compressed ; maxillary exposed ; spinous dorsal 
longer than the soft ; anal with 3 spines, similar to the soft dorsal ; each pelvic 
fin of a spine and 5 soft rays or reduced to a spine only ; caudal fin present. 
Rays of the spinous dorsal equal in number to the vertebrae below them, each 
interneural usually attached to a neural spine ; rays of soft dorsal and anal 
more crowded (except the isolated finlets, when present), about twice as numerous 
as the corresponding vertebrae ; pelvic bones separate, anteriorly extending 
forward to the cleithra and firmly imbedded in the ligament between them. 
Vertebrae 31(15 + 16) to 53(28 -f 25) ; anterior praecaudals without parapo- 
physes, with sessile ribs ; posterior praecaudals with ribs attached at the 
extremities of closed haemal arches ; epipleurals attached to the centra." 

Closely related to the Scombridae, from which, however, they may be descended 
by more than one line, the Gempylidae are quite a varied group of fishes. As Mrs. 
Grey (1953) notes : 

" There is a puzzling scattering of such characters as the presence of a free 
dagger-shaped spine preceding the anal fin, of dorsal and anal finlets, double 
or single lateral lines ; and the presence, absence, or reduction of ventral fins." 



THE FAMILY TRICHIURIDAE 123 

A goodly proportion of the genera are well-illustrated in the paper by Matsubara 
& Iwai (1952). 

The ancestors of the Trichiuridae must undoubtedly be sought among the Gempy- 
linae (Gempylus Cuvier, Nesiarchus Johnson, Mimasea Kamohara), Gempylidae 
which possess an especially elongate body, the head and trunk in particular being 
reminiscent of those of the Trichiurids although the tail seems greatly telescoped by 
comparison and curiously unfinished. In these three genera alone appear the conical 
cartilaginous processes at the tip of the snout and mandibular symphysis which are 
found in the Aphanopodinae ; their skulls are long and low, without prominent 
crests ; they have, like other Gempylids, the typical Trichiurid dentition with the 
three pairs of prominent premaxillary fangs ; their squamation is, at the most, 
vestigial, leading directly to the naked bodies of the Trichiuridae. 

A single row of teeth is present on the palatine in Nesiarchus (personal observation) 
and in Gempylus (Matsubara & Iwai), though Mimasea is said to have none. 

Of these three Gempyline genera Mimasea (Text-fig. 20) is specialized in having a 
double lateral line and a ventral fin-insertion behind the pectoral base ; primitive in 
that the ventral fin is quite well developed, with five soft rays. Despite low median 
fin-ray counts and presumably low vertebral counts therefore, it does not seem a 
likely ancestor to the Aphanopodine Trichiurids. 

Gempylus (Text-fig. 21) has rather less well-developed ventrals, allied, however, 
to a double lateral line and a series of widely-spaced dorsal and anal finlets. The 
proportion of soft dorsal rays to aggregate vertebrae in this genus is very low (18 : 53) 
in comparison with Diplospinus, the most primitive recent Trichiurid (40 : 58) and, 
since the early history of the Trichiurids appears to show soft dorsal rays multiplying 
much faster than the caudal vertebrae, we may feel that the transition from 53 to 58 
vertebrae represents a smaller change than is likely to admit the necesary concomitant 
structural changes (Table VI). 

Nesiarchus, (Text-fig. 22) however, seems to stand very close to the primitive 
Trichiuridae. It has a total of vertebrae (35) near to the minimum of its family 
(31 in Epinnula), allied to a higher number of soft dorsal rays than in Gempylus 
(21-23 : 18) and without detached finlets. The ventral fins are inserted on the per- 
pendicular through the posterior end of the pectoral base and consist each of a spine 
with four smaller soft rays. The skull is well figured by Steindachner (1867) ; apart 
from a broad general resemblance to the skulls of the Aphanopodinae there is a 
striking similarity in the deep opercular notch, nowhere as marked in the other 
Gempylidae and found in only one Trichiurid — the primitive Diplospinus. The 
post-anal spines appear superficially "wrong" ; the first is much larger. But internally 
there is a rudiment of yet another before the first ; there are three well-developed 
spines in Epinnula and it becomes evident that of these the first is to become the 
minute spinule of Nesiarchus and the Trichiuridae (i), the second will become the 
larger spine of Nesiarchus and the principal spine or scute of the Trichiuridae (I), 
and the third, though disappearing, is to signify its claim by a space in the anal 
fin and will contribute to the compound and reinforced anterior basal structure 
whenever this is developed. 



I2 4 



THE FAMILY TRICHIURIDAE 





THE FAMILY TRICHIURIDAE 125 

The body of Nesiarchus is quite naked ; the lateral line is single and descends 
gently to a mid-lateral course along the caudal. The number of pyloric caeca (7) 
is similar to that in the Aphanopodinae. 

Nesiarchus differs from Diplospinus in the lower meristic counts, in having the 
maxillary exposed, in having barbs on the teeth confined to the premaxillary fangs 
and in the external (though not in the internal) structure of the spinous anal fin. 
But the indications from a study of the subsequent evolution of the Trichiuridae 
are that during the addition of 18 vertebrae Nesiarchus would have had plenty of 
time to undergo the modifications needed to produce a Diplospinus. This is the 
view expressed in Text-fig. 23. 

Nesiarchus and Diplospinus therefore may be regarded as the approaches to the 
Gempylid-Trichiurid bridge. Whether the Trichiurinae crossed by the same bridge 
or by a parallel bridge further downstream is still debatable. It is tempting to regard 
the low lateral line in the Trichiurinae as representing the lower limb of the fork 
in another Gempylid ancestor ; but unfortunately, although Trichiurus has a 
longitudinal groove which would serve for an upper limb, no recent Gempylid has a 
lower limb which falls in quite the same way. If, however, the " toothless " palatines 
in Mimasea should, on further examination, prove to be provided with a villiform 
band of teeth, the discovery would be a significant indication of a possible relationship 
and therefore of a diphyletic descent of the Trichiuridae. In this connexion it is 
interesting to observe that the concave lower hind margin of the operculum, charac- 
teristic of the Trichiurinae (though not of Nesiarchus, the Aphanopodinae or the 
Lepidopodinae), makes sporadic appearances among the primitive Gempylidae in 
Epinnula and Neoepinnula. 

Evolutionary trends in the Trichiuridae 

Evolution in the Trichiuridae has resulted from the action, at various rates, of 
the following trends : 

(1) Elongation of the caudal region of the body, least in the stem-forms at any 
level [Diplospinus, Lepidopus) and greatest in the most divergent side-shoots 
(Benthodesmus, Assurger, Tentoriceps). 

(2) Multiplication of the soft dorsal and anal rays, initially at a greater rate than 
that of the adjacent vertebrae. This development, already incipient throughout 
the Gempylidae, is seen proceeding at its greatest rate in Diplospinus and is practically 
in harmony with the multiplying vertebrae in the other Aphanopodinae. 

(3) Multiplication of the caudal vertebrae until eventually (except at the caudal 
tip) each vertebra has one corresponding soft dorsal and anal ray with their associated 
basal elements. This process is nearly complete in Aphanopus and Benthodesmus, 
in which, however, there are usually a very few rays, mainly towards the beginning 
and end of the soft fins, which are not directly related to vertebrae. Except possibly 
in " Lepidopus xantusi " further additions of vertebrae and fin rays proceed in 
unison in the Lepidopodinae and Trichiurinae. 

(4) A slower increase in the number of trunk vertebrae. (See Table VI in conjunction 
with Text-fig. 23). 



126 



THE FAMILY TRICHIURIDAE 



leawii 



LEPIDOPODIN/C 



Assi 



urau* 



Bnvthcd/ismas 



Eiwliuroorammas 




Aphanopus 
APHANOPODlNyfc jy,^*, 



Ntsiatdtus 

GEMPYUNA 



Gempms 
Mmasea, 



Fig. 23. — Suggested relationships of the genera of the Gempylid subfamily Gempylinae 
and of the subfamilies and genera of the family Trichiuridae. 



THE FAMILY TRICHIURIDAE 



127 



Gempylidae 
Gempylinae : 

Nesiarchus nasutus 
Mimasea taeniosoma 
Gempylus serpens 

TRICHIURIDAE 

Aphanopodinae : 

Diplospinus multistriatus 
Aphanopus carbo . 
Benthodesmus tenuis 
Benthodesmus simonyi . 

Lepidopodinae : 

"Lepidopus xantusi" 
Lepidopus caudatus 
Assurger anzac 
Tentoriceps cristatus 
Evoxymetopon taeniatus 
Eupleurogrammus intermedins 

■ Eupleurogrammus muticus 

Trichiurinae : 

Trichiurus lepturus 
Leptur acanthus savala . 



Table VI. 

Dorsal. 

M A „ 

f ^ 

Spines . Soft rays . 



19-21 21-23 

18 16-18 

29-32 18 



32-33 
38-41 

39-42 
45-46 



40 
53-56 

80-88 
108 



102 



(82) 
9 90-97 

(ca. 120) 
(ca. 120) 
10 77 

3 123-131 

3 143-147 



137 
in 



Vertebrae. 

23+12-35 
53 



34+24=58 

42-44+55-56=98-99 

47-52 +75-80= 123-131 

52-53 +101-103 = 153-156 



41+70-73 = 111-113 



32-35 +125-128 = 157-162 
41+150-151 = 191-192 

39-40 + 123-128=162-168 
32-35+124-130 = 159-162 



Data for Nesiarchus (part), Mimasea & Gempylus taken from 
(!953) ) f° r ''Lepidopus xantusi" from Jordan & McGregor (1899) 

(1916) ; for Tentoriceps cristatus from Klunzinger (1884) ; for Evoxymetopon taeniatus from Gill (1863). 
Remainder original. 



Matsubara & Iwai (1952) and Grey 
for Assurger anzac from Alexander 



(5) Progressive reduction of the number of dorsal spines in the higher forms and 
their replacement by soft rays. 

(6) Backward migration of the ventral fins. In the Gempylidae the pelvic girdle 
is embedded in the ligament between the cleithra, and the primitive position of the 
ventral fin-insertions in the Trichiuridae is likewise closely before or behind the level 
of the pectoral base. In Benthodesmus simonyi they are already further back than 
in B. tenuis, while among the Lepidopodinae the migration continues : an eye-diameter 
behind the pectoral base in Lepidopus and Evoxymetopon, an eye-diameter and a half 
in Assurger and finally five eye-diameters in Eupleurogrammus, but all the while a 
ligamentous connection between the pelvic girdle-rudiment and the symphysis is 
maintained. This situation has been discussed by Regan (1909 : 67). 

(7) Hypertrophy of the dorsal musculature, with consequent elevation of the 
posterior confluence of the frontal ridges of the skull into a distinct sagittal crest 
[Lepidopus, Trichiurus) followed at a later stage by an adjacent elevation of the 
ethmo-frontal region continuing this crest forward along the snout [Evoxymetopon, 
Assurger, Eupleurogrammus, Tentoriceps). 

(8) Increase in the number of pyloric caeca. In the Aphanopodinae, as in the 



i 2 8 THE FAMILY TRICHI URID AE 

Gempylinae, the number lies within the range 6-9 (6-8 in 30 Aphanopus counted) ; 
in Lepidopus it is over 20 and may be much higher in Trichiurus (which needs to be 
studied from fresh material) . 

(9) Reduction of the soft anal fin from before backwards. In Diplospinas the 
soft anal extends nearly to the vent, so also in Benthodesmus tenuis. In Aphanopus 
and in B. simonyi the anterior rays are weak and probably of no functional consequence 
in the fin ; in Lepidopus and others only the last 20 rays or so form the true fin. In 
Leptut 'acanthus and in Trichiurus the whole fin is reduced to a series of minute 
spinules, while in Eupleurogrammus the fin as such has ceased to exist and only the 
basal and interhaemal elements remain, firmly interlocking with the haemal arches 
to form a continuous mid-ventral keel. 

(10) Loss of the caudal fin and hypural bones, independently in Trichiurus, 
Lephit 'acanthus , Eupleurogrammus and Tentoriceps. 

(n) Reduction in the extent of the intermuscular (pleural and epipleural) bones. 
In Diplospinus these long bones are a prominent feature in the skeleton and form a 
complete basket surrounding the abdominal cavity, but in all the other genera the 
space which they contain becomes a much smaller portion of the whole. In Eupleuro- 
grammus a small " basket " supported by 14 rather smaller vertebrae is pushed to 
the anterior end of the trunk and is followed by 18-25 vertebrae without epipleurals. 

Pari-passu with the major trends outlined above come sporadic tendencies, 
repeated at different levels : 

(a) Excessive elongation of the body, a possible symptom of evolutionary inertia 
(Benthodesmus, Assurger, Tentoriceps). 

(b) Hypertrophy of the second anal spine, with correlated condensation of the 
anterior basal and interhaemal elements into an enlarged supporting structure. 
(Aphanopus, Leptur acanthus). 

(c) Reduction of the pelvic girdle and fins to an internal rudiment (Aphanopus) 
or their complete loss (Trichiurus, Leptur acanthus) . 

Classification of the Trichiuridae 

The Aphanopodinae as now recognised comprise Gill's group (Aphanopus) with 
the addition of Benthodesmus and Diplospinus, genera recognized since Gill's time. 
They are forms in which the major changes from the Gempyline condition have 
been accomplished but in which the evolution of the Trichiurid caudal may still be 
seen proceeding. The discriminant characters of the primitive Diplospinus have 
already been noted ; it is a satisfactory ancestral form except possibly in the advanced 
barbing of the teeth, a character which, if not merely adaptive, may indicate an 
affinity with the ancestors of the Trichiurinae rather than with the Nesiarchus- 
Aphanopus line. Aphanopus is a secondarily specialized bathypelagic form having 
an enlarged postanal spine and associated endoskeleton. Benthodesmus is an 
attenuate type which has gone some way with Aphanopus (as evidenced by the 
endoskeleton of the anterior anal fin) and then stopped. B. simonyi, evidently 
derived from B. tenuis, shows several evolutionary trends in action in the same genus. 

The Lepidopodinae are equivalent again to Gill's group (Lepidopus, Evoxymetopon) 
with the addition of Eupleurogrammus (removed from Gill's Lepturinae = 



THE FAMILY TRICHIURIDAE 129 

Trichiurinae) and of other genera subsequently recognised — Assurger, Tentoriceps. 
Lepidopus (as represented by L. caudatus) shows a great reduction in the spinous 
dorsal and the early stages in the uplift of the cranial crest and in the backward 
progress of the ventral fins ; at the same time it has attained equilibrium in the 
development of vertebrae and soft fin-rays, and is well on the way towards losing its 
anal fin. The so-called " Lepidopus xantusi " of unhappy memory is inadequately 
known, but would appear to be more primitive than L. caudatus and may even 
deserve generic status in a position between Lepidopus and Diplospinus in the main 
stem. In my opinion L. caudatus represents the termination of a very old line and 
its close similarity of skull to Trichiurus is the result of parallelism and not of any 
closer relationship. The remaining Lepidopodine genera — Evoxymetopon, Assurger, 
Tentoriceps, Eupleurogrammus — have in common an elevation of the ethmo-frontal 
region to continue the sagittal crest forward from the nape to the snout ; in Eupleuro- 
grammus, the only one of this quartet which I have been able to handle, the homo- 
logies in relation to Lepidopus are easily discernible and, together with published 
figures, give sufficient indication of the likely condition in the other three. Evoxy- 
metopon is probably the most primitive of this group, in its shorter body and lower 
median fin-ray counts and in the position of the ventrals and presence of a caudal 
fin, but has a rather steep profile. The ecaudate and highly perfected Eupleurogrammus 
may have been descended from this line, sharing with the Lepidopodines (and 
not with Trichiurus, with which it was formerly classified) the uniseriate palatine 
teeth, median lateral line, ethmo-frontal elevation, ventral fins and rounded 
operculum. The elongate, caudate Assurger and the ecaudate Tentoriceps form 
another like pair. 

The Trichiurinae are now restricted to Trichiurus and Leptur acanthus, the latter 
Fowler's sub-genus upgraded to full generic rank. They are unique among the 
Trichiuridae, not for their loss of a tail (which has occurred elsewhere and indepen- 
dently), but in having a band of villiform teeth on each palatine rather than a single 
series, in having lost the last vestige of a pelvic girdle and fins and in having a low- 
descending lateral line. Other differences assume greater significance in relation to 
these. It is therefore likely that the fundamental cleavage between the Trichiurinae 
and the other two sub-families goes deeper than has previously been supposed. 

It is interesting to observe, in conclusion, that although there has been such a 
great reduction in the number of nominal species formerly placed in Trichiurus the 
residue are now distributed through five genera — Lepidopus, Trichiurus, Leptura- 
canthus, Eupleurogrammus and Tentoriceps. 



REFERENCES 
(Excluding those given under synonymies of genera and species.) 

Berg, L. S. 1940. Classification of fishes, both recent and fossil. Trav. Inst. Zool. Acad. 

Sci. U.R.S.S. 5 : 87-517, figs. 
Boulenger, G. A. 1904. Teleostei. Cambridge Natural History 7 : 539-760, figs. London. 
Capello, F. de B. 1868. Catalogo dos Peixes de Portugal que existem no Museu de Lisboa. 

/. Sci. Math. Phys. Lisboa 1 : 233-264, pi. 



i 3 o THE FAMILY TRICHIURIDAE 

Cuvier, G. & Valenciennes, A. 183 1. Histoire Naturelle des Poissons 8 : 1-509. Paris. 
Gill, T. 1863. Synopsis of the family of Lepturoids, and description of a remarkable new 

generic type. Proc. Acad. nat. Sci. Philadelphia, 1863 : 224-229. 
Goode, G. W. & Bean, T. A. 1895. Oceanic Ichthyology. Spec. Bull. U.S. nat. Mus. 2 : 

1-553, appendix, figs., pis. 
Goodrich, E. S. 1909. Vertebrata Craniata : Cyclostomes and Fishes. In A Treatise of 

Zoology, ed. Ray Lankester, 9 : xvi +518, figs. 
Gregory, W. K. 1933. Fish skulls : A study of the evolution of natural mechanisms. Trans. 

Amer. Phil Soc. 23 : vii + 75-481, figs. 
Grey, Mrs. M. 1953. Fishes of the Family Gempylidae, with records of Nesiarchus and 

Epinnula from the Western Atlantic and descriptions of two new subspecies of Epinnula 

orientalis. Copeia, 1953 : 135-141. 
Gunther, A. i860. Catalogue of the Acanthopterygian Fishes in the collection of the British 

Museum, 2 : 1-548. London. 
Johnson, J. Y. 1865. Description of a new genus of Trichiuroid Fishes obtained at Madeira 

with remarks on the Genus Dicrotus Gunther and on some allied genera of Trichiuridae. 

Proc. zool. Soc. Lond. 1865 : 434-437. 
Jordan, D. S. 1923. A classification of Fishes, including Families and Genera as far as 

known. Stanford Univ. Publ. Biol. Ser. 3 No. 2 : 1-243, i-x. 
Matsubara, K. & Iwai, T. 1952. Studies on some Japanese fishes of the Family Gempylidae. 

Pacif Sci. 6 : 193-212, figs. 
Roule, L. 1927. Considerations sur deux especes abyssales entrees au Musee Oceanographique 

et sur la valeur teratologique possible de l'une d'elles. Bull. inst. Ocean. Monaco, 497 : 

1-12, figs. 
Regan, C. T. 1909. On the anatomy and classification of the Scombroid Fishes. Ann. Mag. 

nat. Hist. (8) 3 : 66-75, figs. 
Starks, E. C. 191 1. Osteology of certain Scombroid Fishes. The Osteological characters 

of the Scombroid fishes of the Families Gempylidae, Lepidopidae and Trichiuridae. Stan- 
ford Univ. Publ. No. 5 : 17-26, pis. 
Steindachner, F. 1867. Ichthyologischer Bericht fiber eine nach Spanien und Portugal 

unternommene Reise. IV : Ubersicht der Meeresfische an den Kiisten Spaniens und 

Portugals. Sitzb. K. Acad. Wien. 56 : 1-106, pis. 
Swainson, W. 1839. The Natural History of Fishes, Amphibians and Reptiles, 2 : 1-452, figs. 

(in Lardner's Cabinet Cyclopaedia, London). 
Tucker, D. W. 1953. The Fishes of the Genus Benthodesmus (Family Trichiuridae). Proc. 

zool. Soc. Lond. 123 : 1 71-197, pis., text-figs. 





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VARIATION, RELATIONSHIPS AND 

EVOLUTION IN THE 

PACHYCEPHALA PECTORALIS 

SUPERSPECIES 

(AVES, MUSCICAPIDAE) 



IAN C. J. GALBRAITH 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 4 No. 4 

LONDON : 1956 



VARIATION, RELATIONSHIPS AND EVOLUTION 
IN THE PACHYCEPHALA PECTORALIS 
SUPERSPECIES (AVES, MUSCICAPIDAE) 




BY 

IAN C. J. GALBRAITH 



Pp. 131-222 ; 8 Text-figures. 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 4 No. 4 

LONDON: 1956 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 4, No. 4 of the Zoological series. 



PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued September, 1956 Price One Pound Four Shillings 



VARIATION, RELATIONSHIPS AND EVOLUTION 
IN THE PACHYCEPHALA PECTORALIS 
SUPERSPECIES (AVES, MUSCICAPIDAE) 

By IAN C. J. GALBRAITH 

SYNOPSIS 

The Australasian species Pachycephala pectoralis is remarkable for the great number and 
variety of geographical representatives which, since they intergrade, must be included in it. 
Although cited in recent evolutionary literature (Meise 1936, Dobzhansky 1937, Mayr 1942, 
Ripley 1945, Cain 1954a), the species has not previously been revised as a whole. It presents 
a wealth of geographically- variable plumage characters, whose relative systematic importance 
can be assessed from their co- variation. The complicated character- geography of the P. 
pectoralis superspecies is here interpreted in terms of colonizations by two major stocks, followed 
by divergence in isolation, great expansions of range, and extensive secondary intergradation. 
Whether two forms, on meeting, will interbreed or coexist as distinct species, seems to depend 
less on their degree of relationship than on internal and external ecological factors. 

CONTENTS 

Page 
Introduction ........... 135 

Scope and presentation . . . . . . . . .136 

Material ........... 137 

Plumage Patterns . . . . . . . . '. ? 137 

Pigments ........... 137 

Patterns . . . . . . . . . . .138 

Retarded and juvenile plumages . . . . . . .139 

Visual significance . . . . . . . . .139 

Variation ........... 140 

Size variation .......... 140 

Male character-geography . . . . . . . .142 

Female character-geography . . . . . . . .145 

Natural Groups .......... 148 

Co-variation and character-complexes . . . . . . 1 48 

Intergradation . . . . . . . . . .149 

Subspecies-groups . . . . . . . . . .149 

P. schlegelii . . . . . . . . . . .150 

Lesser Sundan group A . . . . . . . .150 

Moluccan group B ......... 151 

Solomons group C . . . . . . . . . . 151 

Fijian group D ......... . 153 

P.flavifrons ........... 154 

P. soror ........... 154 

Northern Australian group E . . . . . , .154 

Southern Australian group F. . . . . . . .156 

Southern Melanesian group G . . . . . . . .157 

Widespread group H . . . . . . . .158 

Zool. 4, 4. 10 



134 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

CONTENTS— cont. Page 

Intergradation and Bridgeless Gaps . . . . . .160 

Gene-exchange between subspecies-groups . . . . .160 

Marks of hybridity ......... 165 

Sympatry ........... 166 

Group Affinities . . . . . . . . . .168 

The schlegelii assemblage . . . . . . . .168 

The soror assemblage . . . . . . . . .170 

Arrangement . . . . . . . . . . 171 

The superspecies . . . . . . . . . -171 

Species ........... 172 

Subspecies-groups . . . . . . . . . .175 

Subspecies ........... 175 

Evolution ............ 177 

Geographical speciation in New Guinea . . . . . 177 

Origin of the dichotomy . . . . . . . .178 

Development of isolating mechanisms . . . . . 179 

A suggested course of events . . . . . . . .181 

Rate of divergence ......... 182 

Unexpected uniformity . . . . . . . . .183 

Unexpected diversity . . . . . . . . .184 

Adaptation ........... 186 

Character and climate . . . . . . . . .186 

Character and habitat . . . . . . . . .188 

Variation in habitat . . . . . . . . .189 

Variation in bill size . . . . . . . . .191 

Conclusions ........... 191 

Summary ............ 194 

APPENDIX 

Checklist ............ 195 

Subspecies to be recognized ........ 204 

Notes ............ 205 

P. schlegelii viridipectus ......... 205 

P. pectoralis jubilarii ......... 206 

P. pectoralis atromaculata ........ 206 

P. pectoralis gilolonis .*....... 206 

P. pectoralis ambigua ......... 206 

Female of bynoei .......... 206 

Intergradation of spinicauda with dahli ...... 206 

The subspecies on Teste Island . . . . . . .206 

P. pectoralis fergussonis . . . . ... . . . 207 

P. salomonis ........... 207 

White-throated Pachycephala in the Solomons ..... 208 

P. pectoralis brunneipectus, banksiana and efatensis .... 208 

P. macrorhyncha alfurorum . . . . . . . .208 

P. collaris ........... 208 

P. pectoralis misimae ......... 209 

Saxicola merula .......... 209 

Acknowledgements .......... 209 

Measurements ........... 210 

Weight 211 

Wing and tail lengths . . . . . . . . .211 

Tarsus and culmen lengths, and bill depth . . . . .211 

References ........... 220 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 135 

FIGURES 

Page 

1. Standard plumage patterns . .. . . . . .138 

2. Extreme forms of P. pectoralis . . . . . . . .161 

3. Sympatric species in New Guinea . . . . . . .167 

4. Diagram of relationships . . . . . . . . .169 

5. P. flavifrons, P. p. melanops and P. [p.] caledonica . . . 173 

6. Sketch-map of distribution ....... end-fold 

7. Diagram of colonizations ........ end-fold 

8. Diagram of secondary intergradation ...... end-fold 

TABLES 

Page 

I. Composition of plumage colours . . . . . . .137 

II. Colour-phases in P. flavifrons . . . . . . . .154 

III. Characters uniting the schlegelii assemblage . . . . .170 

IV. Characters uniting the soror assemblage (except subspecies-group H) . 171 

INTRODUCTION 

Pachycephala pectoralis, the Golden Whistler, is probably unique in the richness of 
its geographical variation. More than seventy subspecies can be recognized, extending 
from Java and the Moluccas to Tasmania and Tonga. Many of these are so unlike that 
they would certainly be considered as distinct species, were it not for the more or 
less complete intergradation between them. 

There has been no comprehensive checklist of the species since that proposed by 
Mathews (1930) , in which forms now generally recognized as subspecies of P. pectoralis 
are separated into eleven species. But as early as 1908 Rothschild and Hartert had 
suggested that the distinctive races of the Solomons are conspecific with more 
characteristic P. pectoralis, and this was confirmed twenty years later (Hartert, 1929) 
by the discovery of hybrid populations. Rensch (1931) included the Sumban race 
and its relatives, but not those of the northern Moluccas. Mayr (1932a, b) placed 
even the aberrant forms of northern Fiji in P. pectoralis, because they are connected 
with that species by intermediate forms. The conspecificity of diverse forms has been 
accepted in subsequent lists (van Bemmel, 1948 ; Mayr, 1941a, 1945, 1954a, 1955), 
which together cover almost the whole range of the species. All these forms are 
accepted here as belonging to P. pectoralis. 

P. soror in the hill forest of New Guinea is very like nearby races of P. pectoralis. 
Over most of its range it replaces this species ; but a race of P. pectoralis lives so 
close to populations of P. soror (Rand, 1940) that the barriers between them are 
probably intrinsic (p. 166). Thus two very similar forms seem to be genetically sym- 
patric (Cain, 1953), and must be considered as distinct species ; while others which 
differ much more, and seem to be actually less closely related, intergrade and must be 
considered conspecific. This situation is not unknown in other animals (e.g. Acanthiza, 
Mayr, 1942, 174), but the striking example in Pachycephala has been pointed out 
only by Cain (1954a) as a result of the present review. 



136 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

Scope and presentation 

Four species are considered, which form a single superspecies with a triplet (see 
p. 172) in New Guinea: 

P. schlegelii Schlegel, New Guinea mountain forest from 4,000 or 5,000 to 12,000 ft. 

P. soror Sclater, New Guinea hill forest from 2,200 to 5,200 ft. 

P. pectoralis (Latham), Lesser Sunda Isles and Moluccas to Tasmania and Tonga; 
but absent from New Guinea except for the south-east coast, and disturbed 
habitats in the Snow Mountains between 5,200 and 8,000 ft. 

P.flavifrons (Peale), Samoa. 

No linear arrangement can be satisfactory, since P. pectoralis connects the other 
three species. 

Text-figure 6 (end-fold) gives the ranges of the species and subspecies, and of the 
subspecies-groups of P. pectoralis. The latter seem to be natural groups, though 
because of gene-interchange their boundaries are not sharp and have to be shown 
rather arbitrarily (see Text-fig. 8) : 

Lesser Sundan subspecies-group A (subspecies 1-5). 

Moluccan subspecies-group B (subspecies 6-8) . 

Solomons subspecies-group C (subspecies 9-17). 

Fijian subspecies-group D (subspecies 18-21). 

Northern Australian subspecies-group E (subspecies 22-27). 

Southern Australian subspecies-group F (subspecies 28-33). 

Southern Melanesian subspecies-group G (subspecies 34-38). 

Widespread subspecies-group H (subspecies 39-57). 
P. schlegelii and P. soror have three subspecies each, while P. flavifrons is mono- 
typic. Many forms recognized as distinct subspecies by recent authors have been 
combined in this presentation (see p. 175), although a number of these are distinct 
enough to be separated according to current usage (list on p. 205). 

The use of subspecies names is not helpful to the reader unless he is already 
familiar with the group under discussion. Nor is it usually necessary, since most 
subspecies are easily characterized by their geographical ranges (cf. Wilson & 
Browne, 1953) . In this paper the range citation, given in a condensed and approximate 
form, is followed by a cipher for direct reference to the map (Text-fig. 6). This 
cipher consists of the number of the subspecies within its species, preceded by the 
subspecies-group letter for subspecies of P. pectoralis. Where infrasubspecific 
variation is discussed, parts of the subspecies range are indicated by lower-case 
suffixes. Thus "Sumbawa to Alor A3" indicates the form of P. pectoralis on Sumbawa, 
Flores, Lomblen, Pantar and Alor (which from "Lomblen to Alor A3b". is slightly 
larger and larger-billed). The exact range of any form can be found from the check- 
list (p. 195), which is lettered and numbered to correspond. 

Subspecies names are useful in referring to forms whose geographical ranges are 
diffuse or difficult to define: " dahli E25 " is used for the subspecies of P. pectoralis 
which ranges from south-eastern New Guinea and Fergusson Island to many small 
islands in the Bismarck Archipelago. The range of dahli is shown inset on the map. 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 



137 



Material 

I have seen at least one adult male and one adult female of every subspecies 
recognized by recent authors, except for the following : no specimens of H46 and 
H50 ; no adult male of D21; no adult female of C15, G37a & b, H49, H52 and H56 
(juveniles seen) and E25C (female unknown). I have examined the following types in 
the British Museum (Natural History) : fulvotincta Wallace A3a, mentalis Wallace 
B6, neglecta Layard (= Di8b), aurantiiventris Seebohm Diga, torquata Layard D2oa, 
fuliginosa Vigors & Horsfield F28b, fusca Vigors & Horsfield (= F3ob), variegata 
Gray (= G34), cucullata Gray G36, chlorurus Gray G37a, intacta Sharpe G37d, 
fuscoflava Sclater H42, xanthocnemis Gray (= H43b), clio Wallace H45, collaris 
Ramsay H47a, vitiensis Gray H55, klossi Ogilvie-Grant and bartoni Ogilvie-Grant 
(P. soror 2 & 3). 

Unfortunately, many of the available series were very short, and the measurement 
tables compiled (p. 212) are inadequate for proper statistical treatment. Individual 
variation and fine geographical variation are therefore not considered in this paper. 
Where the available material was inadequate full use has been made of published 
descriptions and measurements, especially those of Mayr (1932a, b, and 1954a). 

I have studied P. pectoralis in the field on Guadalcanal Cn and San Cristobal Ci7a, 
and seen and heard it near Sydney F3ob, on Lord Howe and Norfolk Islands F32 
& 33, and on Efate and Santo G37b & d. I have also studied P. implicata (which is 
rather closely related to the pectoralis superspecies) in the mountains of Guadalcanal 
(Cain & Galbraith, 1956). 

PLUMAGE PATTERNS 

Pigments 

All the colours in the various plumages of the superspecies are produced by 
combinations of yellow, black and brownish pigments (Table I). The yellow pigment 
is soluble in boiling alcohol or pyridine, and turns a transient blue-green with con- 
centrated sulphuric acid. It is therefore a carotenoid (Cain, 1950, 104). Carotenoid 



Table I. — Composition of Plumage Colours. 



Melanins. 


None. 


None ..... 


White 


Eumelanin : 




Barbs clear, barbules satu- 


Grey 


rated 




Granules in barbs, barbules 


Dull black 


saturated 




Saturated .... 


Black 


Phaeomelanins : 




Pale 


Sandy to 




dull pink 


Deep .... 


Brown to 




rufous 



Carotenoid. 
a 



V. pale. Pale. Deep. 

Pale yellow Lemon -yellow Golden- to 

orange-yellow 



Olive-grey 



Olive-green Golden-olive 
Olive-black — 



Cream 



Russet 



Cinnamon 

to vinous 

Tawny-orange 



Citrine 



138 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

tends to be more concentrated in the barbs than in the barbules of the feather. The 
other pigments, if not saturated in both elements of the vane, are more concentrated 
in the barbules than in the barbs. They are granular. Under the microscope black 
granules still appear intense black, while brown ones vary considerably in hue and 
intensity. The black and brown pigments are presumably eumelanins and phaeome- 
lanins. Parts of the vane devoid of melanins are filled with minute bubbles, which by 
multiple internal reflection add to the brilliance of the whites and yellows. Most 
contour feathers are grey basally, with the black granules forming bands across the 
barbules. 

Patterns 

It is convenient to describe the variation of the male pattern throughout the 
superspecies in terms of departure from a standard. The descriptions will be briefest, 
and the peculiarites of the various forms most clearly apparent, if the pattern chosen 
as standard combines all the variants which are more common than their alternatives. 
This condition is fulfilled by the pattern of males of P. pectoralis in the Bismarcks 
H48-51, which is shown diagrammatically in Text-fig. 1 (top), and described below. 




Fig. 1. — Diagram of the standard plumage patterns for the superspecies. Male (above, 
facing left) H48-51 ; female (below, facing right) H52. Not to scale. Colour key : 
diagonal hatching, grey (without stipple) or olive (with stipple) ; cross-hatching, brown 
and rufous hues ; stipple, yellow ; black, black ; white, white. The same conventions 
are used in all the diagrams of plumage pattern (Text-figs. 1-3 and 5). 

Adult male. White chin and throat separated by black gorget, between auriculars, 
from golden-yellow underparts; fore-edge of gorget formed by black tips of white 
feathers, hind-edge by black feathers overlying yellow ; black cap and auriculars 
separated by yellow collar across hind-neck from golden-olive mantle ; wings dull 
black with olive outer edges to all feathers, fading to grey towards tips of outer 
primaries ; inner edges of quills whitish ; tail and upper tail-coverts black with olive 
tips, and olivaceous traces at bases of outer quills. 

It is harder to select a standard for the female plumage, since the patterns are 
much vaguer than those of the males, and the variation is more quantitative than 
qualitative. The following description and the diagram (Text-fig. 1, bottom) best 
fit some females of the widespread group H, and especially those from Ndeni H52. 

Adult female. Throat pale buff with faint dark fringes at sides ; gorget brownish, 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 139 

vaguely defined, grading into brownish wash on breast and flanks ; mid-belly and 
under tail-coverts free from melanins ; under tail-coverts pale yellow, breast and 
belly washed with yellow ; upper parts citrine ; collar, rump and tail brighter ; cap 
greyer than mantle, auriculars cinnamon ; wings blackish-brown with citrine outer 
edges to all feathers, fading to greyish towards tips of outer primaries ; inner edges 
of quills whitish. 

Retarded and juvenile plumages 

Males are sometimes found in plumage like that of the adult females (" II Phase " 
of Mayr, 19320, b), though commonly with more intense carotenoid. They agree with 
adults in tail shape and in having black bills, and their testes are often enlarged 
(Baker, Marshall & Harrison, 1940). They are evidently adult males in a " retarded " 
plumage (Mayr, 1933) : in Victoria and Tasmania at least they are known to breed 
in this condition (Chandler, 1912 ; Howe, 1927 ; Lawrence, 1952). 

Juveniles of both sexes have brown or straw-coloured bills, and softer wings and 
more pointed tail-feathers than adults. In pattern they resemble adult females, 
except that they have the edges of the wing feathers rufous, the cap olivaceous, the 
mantle often browner and the carotenoid pigment more dilute (I Phase). Nestlings 
are commonly rufous all over ; but in the lesser Sunda Isles (nestling of A3b seen) 
are greyish with darker shaft-streaks beneath. Only Mayr (1932a, b) has systemati- 
cally described immature plumages, and I have not examined sufficient immature 
skins from parts of the range not treated by him to attempt a survey of their varia- 
tion. 

At first sight the female and juvenile patterns seem very different from that of 
the adult male. But the distribution of pigments is much the same — except that 
carotenoids are greatly diluted, there are no solid blacks, and eumelanins are more or 
less replaced by phaeomelanins. As a result, these patterns are less sharply defined 
and stereotyped than that of the male, and show a greater range of colours. 

Visual significance 

In many birds the upperparts are cryptic, while the underparts bear a conspicuous 
pattern which plays a part in intraspecific display. The standard male pattern of 
P. pectoralis is a good example of this. Both the ventral and the dorsal pattern 
embody transverse bands which contrast strongly with the ground-colour (see Cott, 
1940) . The bright underparts contrast with the forest background, so that the black 
gorget (exploiting maximal tone-contrast against the white, and an especially effective 
colour-contrast against the yellow) is an emphatic feature. The olive upperparts, on 
the other hand, conform with the background, and the yellow collar is disruptive in 
effect. Females and juveniles tend to be inconspicuously coloured, with neither 
emphatic nor markedly disruptive features. 

This common species has been largely neglected by Australian ornithologists 
(except Lawrence, 1952), and nothing seems to be known of the part played in 
intraspecific display by the conspicuous male ventral pattern. Although we saw 
neither courtship nor aggressive displays in the Solomons, the general behaviour of 



i 4 o THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

these birds (Cain and Galbraith, 1956) is much what might be expected from a study 
of their colour-patterns. The males show themselves freely to a ground observer as 
they move about below the canopy, whistling loudly. But although they will sing 
from bare branches overhung by foliage, they avoid all perches exposed to the sky. 
Females of P. pectoralis on Guadalcanal Cn, like those of many other subspecies, 
are dull in colour. They are very silent and skulking, seldom seen as they move 
about in dense thickets near the ground. Those on San Cristobal C17 are as bright 
yellow beneath as the males, but lack the black gorget. They are often seen in company 
with the males in the upper substage, usually silent, but occasionally heard to give 
alarm calls and quiet whistles. A related species in the mountains of Guadalcanal 
(P. implicata) has the females actually more conspicuous than the males. They are 
not more retiring than the males, and are more vocal than the females of P. pectoralis. 
The males of P. implicata are inconspicuously coloured in olive and blackish, and are 
much more silent and retiring than those of P. pectoralis. On the other hand, a pair 
of this species was seen feeding exposed in the tops of small trees: perhaps the direction 
of predation is less important to these more uniformly-coloured birds. 



VARIATION 

Geographical variation in the superspecies is conspicuous and complex. Marked 
sexual dimorphism is normal, but in three widely-separated races of P. pectoralis 
the males are permanently hen-feathered, while P. flavifrons has cock-feathered 
females. The sporadic occurrence, in sexually dimorphic forms, of breeding males in 
female plumage shows that hen-feathering is not very important here, either as an 
indicator of genetical change or as a potential barrier to interbreeding. 

The bold pattern of the male is subject to striking variation. For example, in 
some forms the throat is yellow instead of white, and some of these lack the black 
gorget. These differences are not unifactorial, since for almost every pair of contrasted 
characters intermediate states are to be found. Single characters cannot be used to 
delimit natural groups, since not one variant is confined to a single subspecies-group 
of P. pectoralis (as distinguished by the whole constellation of characters of both 
sexes) and shown by every race belonging to that group. Most are both polyphyletic 
in origin and labile within groups. The marked geographical variation of the female 
pattern does not present the same appearance of discontinuity, and there is less 
temptation to erect a classification based on single characters of the females. 

Size variation 

The largest subspecies of P. pectoralis have wings over 30% longer than the smallest. 
If their proportions are the same, they should therefore be more than twice as heavy. 
Unfortunately, weight measurements are available for very few forms in the super- 
species, so that wing lengths are for the present the best indication of body size. 
Certain subspecies which are evidently very closely related differ markedly in size, 
which is therefore not generally of much importance in assessing relationships ; but 
some of the discontinuities between subspecies-groups are marked by sharp changes 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 141 

in size as well as in other characters. A coarse grading into size-classes adequately 
expresses the major variation, which more detailed treatment would tend to obscure. 
In any case, too few specimens of many forms were available for quantitative treat- 
ment to be satisfactory. Snow (1954) has pointed out that quantitative differences 
(in size, proportions, intensity of pigmentation, etc.) are often adaptive, and cannot 
be used as clues to relationships and past history until their present environmental 
relations have been worked out. Conversely, where the past history has been as com- 
plex as in the pectoralis superspecies, such differences cannot be shown to be 
adaptive until relationships have been established (by studying the co-variation 
of more stable characters). Two forms on neighbouring islands might differ in size, 
not because the climates were different but because they were the same, if the 
forms belonged to different subspecies-groups whose size /temperature relationships 
(Snow, 1954, 22) had been evolved under different climates. 

The various forms are graded into five classes on the average wing lengths of 
adult males. But the arbitrarily-selected limits of these classes have in a few cases 
been relaxed, to avoid separating closely-related forms whose small difference in 
size happens to transgress them. The southernmost Australian forms are classed as 
" large ", not as " very large ", because the appearance of the skins (prepared by 
many different collectors) strongly suggests that they are considerably smaller birds, 
though with relatively long wings (perhaps in relation to their nomadic habit) than 
the " very large " ones of the Solomons and elsewhere. This can only be decided 
when weight measurements are available. The middle class, containing the most 
forms, is of wing lengths from 90 to 95 mm.; those from 80 to 85 mm. are graded as 
" very small", from 85 to 90 as " small ", from 95 to 100 as " large " and over 100 as 
" very large ". The following forms fall outside the middle class : 

Very small birds. Sumba, Java to Alor, and Flores Sea and Salayer A1-5 ; 
midwestern Australia E22 a-c (size increasing north-eastwards) . 

Small birds. Rennell and San Cristobal C16-17 ; northern Australia to southern 
New Guinea E22d-24 ; New Caledonia, New Hebrides and Vanikoro G34 & 36-38 ; 
Timor to Damar H39-41 ; Sula and Peleng Isles H45-46 ; large islands in the 
Bismarcks H48 ; Kandavu H54 ; P. soror ; P. schlegelii ; P. flavifrons. 

Large birds. Ternate, Tidore and Obi B7-8 ; Vella Lavella to Gatukai C13 & 
15 ; Tasmania and southern Australia F28-30, size decreasing northwards ; Tabar 
H50 ; Lau archipelago H56. 

Very large birds. Bougainville to Malaita, and Rendova and Tetipari C9-12 
& 14 ; Loyalty Isles G35 ; Tenimber Isles H42 ; Lihir H49 ; Tonga H57. 

Differences in bill size are even more marked, with the largest bills some 60% 
longer than the smallest, and the bill may be stubby, stout or slender. However, 
closely related forms differ in bill size even more than in wing length, and the accuracy 
of this measurement is not sufficient for the forms to be graded with any degree of 
certainty from the scanty data available. Tail and tarsus lengths are only comparable 
when expressed as relative lengths, which vary between individuals much more than 
the absolute measurements. Therefore none of these measurements is dealt with 
comprehensively, though there is some discussion of climatic correlation in certain 
parts of the range (p. 186). 



142 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

Male character -geography 

Some of the most conspicuous variations appear to be qualitative and discon- 
tinuous, but hybrid forms often show intermediate states. Probably most of the 
variations are in fact potentially continuous — but it is convenient to deal with the 
geographical distribution of the more important variations in terms of discontinuous 
"characters", at the same time mentioning the occurrence of intermediate conditions. 

(i) Hen-feathered. Rennell C16, Norfolk Island F33 (sexes almost indis- 
tinguishable) ; Salayar A5 (male in a female type of plumage, but more advanced in 
character than its own female). 

The hen-feathered races were considered as separate species until Mayr (1932a, 5) 
pointed out the relative unimportance of the character. Since individual males of 
dimorphic subspecies can breed in female plumage, hen-feathering involves neither a 
radical change in the genome nor complete breakdown of courtship and territorial 
displays. Variations of the male pattern such as "no gorget ", "no collar ", " cap 
pale " and " tail pale " may be considered as partial hen-feathering, and the San 
Cristobal form C17 especially shows several such indications. 

The three fully hen-feathered forms are not considered further in the male character- 
geography. 

(2) Chin black. Sumba Ai ; Morotai to Bat j an B6-7 ; Solomons except San 
Cristobal C9-15 ; P. schlegelii, P. flavifrons. 

(3) Throat and chin black. Tonga H57. 

In P. flavifrons there is much black at the base of the throat feathers, which shows 
as irregular barring. Another approach to the black-throated condition is shown 
in the reduction of the throat patch on Sumba Ai (where black malar feathers 
intervene between it and the auriculars), from Java to Alor A2-3, and in P. soror 
and P. schlegelii. 

(4) Throat yellow. Solomons C9-17 (variable in hybrids on small islands off 
Shortland E26) ; northern and central Fiji D18-21 ; Tenimber Isles H42 ; P. flavi- 
frons (individually variable) . 

Juveniles on Vanikoro G38 have yellow throats, and there is a tendency for the 
throats of juveniles in the New Hebrides G37 and on Utupua H53 to be yellowish 
(Mayr, 19326). A few yellow feathers occasionally appear in the white throats of 
adult males. I have seen these in males of P. pectoralis from Teste Island E25b, 
Lord Howe Island F32, New Caledonia G34, Loyalty Islands G35, Aneitum G36 
and Kandavu H54, and in P. soror from the Snow Mountains 2a. But they seem to 
occur most frequently in P. schlegelii — in eight of thirty-one adult males examined. 

(5) Bases of throat-feathers white. Obi B8 ; Solomons C9-17 ; northern 
and central Fiji D18-21 ; midwestern Australia to small islands in the Bismarcks, 
and hybrids off Shortland E22-26 ; Vanikoro G38 ; Tenimber Isles H42 ; Sula and 
Peleng Isles H45-46 ; Louisiades H47 ; Ndeni, Santa Cruz islets and Utupua H52-53. 

(6) No gorget. Malaita C12; Viti Levu and Vanua Levu Di8a and 19a; 
P. flavifrons. 

The gorget is more or less broadly interrupted in individual males on the islands 
between Koro and Taviuni on the one hand, and Viti Levu and Vanua Levu on the 
other— Di8b, 19b & c. 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 143 

(7) Fore-edge of gorget defined by pale feathers. Sumba and Java to 
Alor A1-3 ; northern Moluccas B6-8 ; Solomons except San Cristobal (and the 
gorgetless forms on Malaita and Rennell) C9-11 & 13-15 ; Ceram H43 ; Ndeni and 
Santa Cruz islets H52 ; P. schlegelii. 

In these forms the black of the gorget deeply undercuts the white or yellow of the 
throat, whereas in the standard pattern the edge of the gorget is defined on the under- 
lying layers of feathers at about the same level. On San Cristobal C17, in the hybrids 
on the outliers of Shortland E26, and on Buru H44, the gorget undercuts the throat- 
patch, but there are also some black tips. 

(8) Gorget cut off from auriculars. Northern Moluccas B6-8 ; New Caledonia 
G34 ; Ceram H43 ; Ndeni and Santa Cruz islets H52. 

In all these forms the throat-feathers are unusually long, extending further back 
than the auriculars. They are best developed from Morotai to Bat j an B6-7 (where 
they cover most of the gorget), and least so on New Caledonia G34 (where only the 
lateral feathers are appreciably lengthened). In B6-7 and G34 the gorget is itself 
reduced, and does not join the auriculars even beneath the white feathers. On Vella 
Lavella and Ganonga C13 the yellow feathers extend behind the auriculars, but the 
gorget is too broad to be cut off. 

(9) Gorget very broad. Vella Lavella and Ganonga C13 ; P. schlegelii. 

(10) Gorget very narrow. New Caledonia G34 ; Kandavu H54 ; P. soror. 

(11) Breast rufous. Sumba Ai (intense, extending to flanks and belly) , Java A2a 
(less intense, especially on belly), Bali A2b (still less intense on flanks and belly), 
Sumbawa to Alor A3 (confined to band behind gorget), islands in Flores Sea A4 (very 
faint, barely detectable on belly and flanks) ; Santa Anna Ci7b and individuals on 
San Cristobal Ci7a (patch behind gorget) ; New Caledonia G34 (patch behind 
gorget) ; P. schlegelii (extending to flanks and belly ; very intense from Snow 
Mountains to south-eastern New Guinea 3, somewhat less so in Vogelkop and Cyclops 
Mountains 1 & 2). 

There is a rufous wash centred on the vent on Taviuni, Koro and Vatu vara 
D20-21, Ceram H43, Ngau H55, and in individuals on Aneitum G36. 

Where very pale rufous phaeomelanin combines with pale yellow carotenoid (as 
in the Flores Sea A4), the tawny-orange produced is rather like the orange-yellow of 
very intense carotenoid (as on Vanua Levu D19). The difference is easily seen 
through the microscope, since the phaeomelanin appears granular. 

(12) Carotenoid very deep (orange- yellow). Vella Lavella and Ganonga 
C13 ; Vanua Levu to Koro and Vatu vara D19-21 ; Louisiades H47 ; Lihir H49 ; 
Manus H51. 

(13) Carotenoid pale (lemon- yellow and greenish-olive). Mid- and north- 
western Australia E22, deepening northwards E22a to 23 ; Tasmania and southern 
Australia F28-30, deepening northwards F29 to 31 ; New Caledonia, New Hebrides 
and Vanikoro G34 & 36-38 ; Babar H40 ; Utupua H53 ; P. soror. 

(14) Forehead yellow. Viti Levu, Vanua Levu and Vatu vara Di8a, 19a & b 
& 21 (sometimes a few yellow feathers on Ovalau, Rambi and Kio Di8b & 19c) ; 
P.flavifrons (sometimes white, in white- throated individuals). 

(15) Cap pale. Santa Anna Ci7b (olive, with blackish lores and auriculars) and 



i 4 4 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

San Cristobal Ciya. (variable, from olive to black) ; New Caledonia G34 (grey) ; 
Aneitum G36 (dull black with more or less olive scalloping) . 

(16) No collar. Vella Lavella and Ganonga C13, Ndeni and Santa Cruz islets 
H52, P.flavifrons (melanic forms with no trace of a collar) ; Fiji, except for Vatu vara 
and Kandavu, D18-20, H55-56 (black of cap extends far down on hind-neck, 
followed by a yellowish trace) ; Malaita C12, New Caledonia G34, P. soy or (yellowish 
trace) ; San Cristobal C17 (yellowish trace on hind-neck, clear yellow patches at 
sides of neck) . 

The collar is very vague and narrow on the islands in the Flores Sea A4, from 
Kulambangra to Gatukai C15 and on the Tenimber Islands H42. It is narrow and 
washed with olive on the nape on Sumba and from Java to Alor A1-3, on Rendova 
and Tetipari C14, in mid- and north-western Australia E22, in Tasmania, southern 
and eastern Australia and on Lord Howe F28-32, from the Loyalty Isles to Vanikoro 
G35-38, on Timor 1139a, the Louisiades H47, Utupua H53 and Kandavu H54. 

(17) Mantle black. Vella Lavella and Ganonga C13, Ndeni and Santa Cruz 
islets H52, P. flavifrons (wholly black) ; islands in Flores Sea A4, Rendova and 
Tetipari C14, Taviuni and Koro D20, Ngau and Lau archipelago H55-56 (feathers 
black-centred, individually variable from olive with concealed black spots to black 
with olive scalloping) . 

(18) Wing black. Islands in Flores Sea A4 ; Vella Lavella, Ganonga, Rendova 
and Tetipari C13-14 ; P. flavifrons ; P. schlegelii. 

In these forms the pale edges to the wing-feathers are absent or exceedingly narrow. 
The pale edges are narrow, and the primary-coverts entirely black, on Taviuni, 
Koro and Vatu vara D20-21, in dahli E25, on Ceram and Bum H43-44, Ndeni and 
Santa Cruz islets H52 and Ngau and the Lau archipelago H55-56. 

(19) Olive very dark. Kulambangra to Gatukai C15 ; Fiji D18-21 & H54-56 ; 
Ceram and Buru H43-44 ; P. soror ; P. schlegelii. 

Normally the bases of the barbules, as well as the barbs, are yellow, giving a 
more or less pronounced herring-bone pattern under the microscope. In the above 
forms the barbules are black down to their junctions with the barbs. 

(20) Olive tinged with brownish. San Cristobal C17 ; Koro and Taviuni 
D20 ; Ceram H43 ; Louisiades H47 ; individuals on Aneitum G36. 

The phaeomelanic wash is deepest on tail, upper tail-coverts, rump and secondaries. 

(21) Wing quills grey-edged. Midwestern Australia to small islands in the 
Bismarcks E22-25 '> Tasmania F29 ; Damar H41 ; P. soror in Snow Mountains 2a. 

The wings are unusually grey in southern Australia F28 & 30, becoming more olive 
northwards ; and in eastern Fiji D 19 & 21 & H56, becoming more olive westwards 
(towards Viti Levu and Kandavu D18 & H54). They are also unusually grey in all 
populations of P. soror. They are variable in colour in the hybrids on outliers of 
Shortland E26. 

(22) Upper wing-coverts yellow-edged. Viti Levu and Vanua Levu D18-19 ; 
midwestern Australia to small islands in the Bismarcks E22-25 ; Tasmania, southern 
and eastern Australia and Lord Howe F28-32 ; Loyalty Islands G35. 

The upper wing-coverts are edged with yellower olive than the mantle in many 
forms ; those listed above show this most conspicuously. 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 



145 



(23) Upper tail-coverts all-olive. Kulambangra to Gatukai C15 ; San 
Cristobal C17 ; northern Fiji, except Koro and Vatu vara, Di8-2oa ; mid- and 
north-western Australia E22 ; Tasmania, southern and eastern Australia and Lord 
Howe F28-32 ; New Caledonia to Banks Islands G34-37 ; Timor H39 ; Tenimber 
Isles H42 ; Louisiades H47 ; Kandavu H54 ; Tonga H57 ; P. soror. 

(24) Tail pale. Tasmania F29, New Caledonia, Loyalty Islands and Aneitum 
G34-36, Timor H39, western Louisiades H47a (no solid black) ; San Cristobal C17 
(more or less black) ; south-western and South Australia F28, New Hebrides from 
Erromango northwards G37, Misima H47b, Kandavu H54, P. soror except the 
Snow Mountains 1 & 3 (black subterminal patch) ; south-eastern Australia and 
Lord Howe F30 & 32, Babar H40, Rossel H47C (more than half black) ; mid- western 
Australia E22a-c, southern Queensland F3ia, Damar H41, P. soror in Snow Moun- 
tains 2 (wide olive edges basally) ; north-western Australia E22d, northern Queensland 
F3ib (narrow olive edges basally). 

The pale part of the tail, usually olive, is grey from south-western Australia to 
Tasmania and Victoria F28~3oa, and sometimes greyish in northern Queensland F3ib. 

(25) Upper tail-coverts all-black. Sumba, Java to Alor, and islands in 
Flores Sea A1-4 ; Morotai to Obi B6-8 ; Vella Lavella and Ganonga C13 ; Ceram, 
Sula and Peleng Islands H43 & 45-46 ; Ndeni and Santa Cruz islets H52 ; P. 
schlegelii ; P. flavifrons. 

Even in these forms, the shortest coverts usually have narrow olive tips. Males on 
Buru H44, and from some other localities, have the tips of all the coverts extremely 
narrow. 

(26) Tail all-black. Sumba Ai ; Morotai to Batjan B6-7 ; Vella Lavella 
and Ganonga C13 ; Ndeni and Santa Cruz islets H52 ; P. schlegelii ; P. flavifrons. 

There are sometimes narrow and obscure olivaceous tips to the lateral tail feathers, 
even in these forms. The tips are unusually narrow from Java to Alor A2-3, on Obi 
B8, in most of the Solomons C9-12 & 14-15, and from Ceram to Peleng H43-46. 



Female character -geography 

(1) Cock-feathered. P. flavifrons (throat and forehead patterns less sharply 
defined than in the male) . 

Certain single characters of the female (such as a white throat, a sharply defined 
gorget, bright yellow underparts, a pure grey cap and pure olive mantle, and a partly 
black tail) tend perhaps towards the male pattern. Often they appear independently, 
but on San Cristobal C17, and also in northern members of group E and in P. soror, 
most of the characters mentioned occur together — these forms have an advanced 
type of female plumage. Individual females of the race on Vella Lavella Ci3a 
(whose males are all-black above) have black gorgets (Mayr, 1932a, 17). A partly 
cock-feathered female from Malaita C12 is reported by Mayr (1932a, 21). I have seen 
several specimens which (though sexed as females) are partly or wholly in the plumage 
of the adult male, but consider it unwise to rely on the sexing of these, since they 
carry no indication that the collector had noticed the discrepancy between gonads 
and plumage. 



i 4 6 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

P. flavifrons is not considered further in the female character-geography. 

(2) Bill pale. Solomons C9-17 (palest from Guadalcanal to Kulambangra 
and on Rennell Cioc-11 & 15-16, darkest on Malaita, Vella Lavella and Ganonga 
C12 & 13) ; Rambi and Kio D19C. 

(3) Throat without melanic wash. Islands in Flores Sea and Salayer A4-5, 
midwestern Australia E22a-c, New Caledonia to Banks Islands G34-37, Timor and 
Babar H39-40, Louisiades H47, Utupua H53, P. soy or (wholly white) ; north- 
western Australia to small islands in the Bismarcks, hybrids near Shortland and the 
Snow Mountains E22d-27, Vanikoro G38, P. schlegelii in the Vogelkop and Cyclops 
Mountains 1 & 2 (white, more or less mottled) ; San Cristobal C17 (pure yellow). 

(4) Throat yellow. Choiseul to Russel Isles, central Solomons, Rennell and San 
Cristobal Cio & 13-17 ; Tenimber Isles H42. 

(5) Cheeks yellower than throat. Bougainville, Guadalcanal and Malaita 
C9, 11 & 12 ; Viti Levu and Vanua Levu Di8a & 19a. 

(6) Throat barred. Morotai to Obi B6-8 ; Bougainville, Guadalcanal and Malaita 
C9, 11 & 12 ; Viti Levu and Vanua Levu Di8a & 19a, and individuals on Ovalau, 
Rambi and Kio Di8b, 19D-C ; north-western Australia to small islands in the 
Bismarcks, hybrids near Shortland, and the Snow Mountains E22d-27 ; Tasmania, 
southern and eastern Australia, Lord Howe and Norfolk Islands F28-33 ; Damar 
H41 ; Sula and Peleng Islands H45-46 ; Tonga H57 ; P. schlegelii (faintly in the 
Vogelkop 1). 

There is no sharp distinction between subterminal barring and terminal fringing 
of the feathers. There is a tendency towards heavy fringing, not entirely confined to 
the sides of the throat, from New Caledonia to the Banks Islands G34-37, in the 
Louisiades and Bismarcks H47-48, and on Utupua H53. 

(7) Throat and breast shaft-streaked. P.p. dahli, hybrids near Shortland, and 
Snow Mountains E25-27 ; Vanikoro G38 ; Manus H51. 

(8) Underparts shaft-streaked. Solomons except San Cristobal C9-16 ; 
Viti Levu and Vanua Levu Di8a & 19a, and individuals on Ovalau, Rambi and Kio 
Di8b & 19D-C. 

(9) Gorget eumelanic. Morotai to Bat j an B6-7, P. schlegelii east of the Vogel- 
kop 2-3 (pure grey) ; Choiseul and Kulambangra to Florida, Rennell and San 
Cristobal Cio & 15-17, Loyalty Islands and Vanikoro G35 & 38 (olive) ; Bougain- 
ville and Malaita C9 & 12, northern Australia E22-23, Tasmania and southern 
Australia F28-30, New Caledonia, and New Hebrides from Erromango northwards 
G34, 37a-c & e (considerably greyer than standard). 

(10) Gorget yellow- washed. Choiseul to Russel Islands, central Solomons, 
Rennell and San Cristobal Cio & 13-17 ; Loyalty Islands and Vanikoro G35 & 38 ; 
Babar and Tenimber Isles H40 & 42 ; Louisiades H47. 

(11) Breast and flanks olive. Morotai to Bat j an B6-7 ; Choiseul and Kulam- 
bangra to Florida, and Rennell Cio & 15-16 (pale) ; Snow Mountains E27 (pale, 
confined to narrow band behind gorget) ; P. soror (pale) ; P. schlegelii (deep in 
Snow Mountains and south-eastern New Guinea). 

Many forms have a combination of carotenoid and phaeomelanins on the under- 
parts, but this ventral olive is rare. 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 147 

(12) Underparts uniform. Koro and Vatu vara D20D-21, and individuals on 
Ovalau and Taviuni Di8b & 20a ; Tenimber Isles H42 ; Mussau and Lihir H48d~49 ; 
southern Fiji H54-56. 

On Ceram and Bum H43-44 and in the Bismarcks H48a-c, the gorget is little 
darker than the throat and mid-belly. 

(13) Belly washed with melanins. Islands in Flores Sea A4 (pinkish) ; 
Bougainville, Guadalcanal and Malaita C9 & n-12 (greyish or rufous buff) ; Rendova 
and Tetipari C14 (deep rufous) ; Fiji D18-21, H54-56 (greyish on Viti Levu and 
Vanua Levu Di8a & 19a, cinnamon on Koro, Vatu vara and in southern Fiji D20D-21 
& H54-56, individually variable between Koro and the large islands Di8b & 19b- 
20a) ; south-western and South Australia F28a-b (pinkish), Tasmania, Victorian 
mallee and northern Queensland F28C, 29 & 31b (buffy) ; New Caledonia G34 
(pale buffy) ; Timor H39 (pinkish) ; Damar, Tenimber Isles, Ceram to Sula Isles, 
Bismarcks except Tabar H41-45 & 48-49 (brownish, more or less mixed with 
yellow). 

(14) Breast and belly deep yellow. Morotai to Obi B6-8 (deepest on Obi 
B8) ; Choiseul and Vella Lavella to Florida, and San Cristobal Cio, 13, 15 & 17 ; 
northern Australia, dahli and Snow Mountains E23, 25 & 27 ; Loyalty Islands and 
Vanikoro G35 & 38 ; Babar H40, Peleng Isles H46, Louisiades H47, Tabar H50, 
Manus H51, Utupua H53, Tonga H57 ; P. soror ; P. schlegelii. 

(15) Breast and belly scarcely yellow. Sumbawa to Alor and islands in 
Flores Sea A3-4 ; Bougainville, Guadalcanal and Malaita C9, 11 & 12 (juveniles 
may be yellower) ; Fiji except Kandavu D 18-21, H55-56 ; midwestern Australia 
E22a-c ; Tasmania and southern Australia F28-30 ; Malekula and Santo G37d ; 
Timor, Damar and Buru H39, 41 & 44. 

(16) Under tail-coverts not yellow. Tasmania and southern Australia 
F28-30. 

Here the female plumage is almost devoid of carotenoid, except for a circlet of 
pale yellow feathers at the vent. 

(17) Cap olive. Solomons C9-17 (may be obscured by heavy rufous wash). 

(18) Cap grey. Salayer A5 ; Morotai to Batjan B6-7 ; midwestern Australia 
to small islands in the Bismarcks E22-25 ; Tasmania, southern and eastern Australia, 
Lord Howe and Norfolk Island F28-33 ; Vanikoro G38 ; Tabar H50, Lau archipelago 
and Tonga H56-57 ; P. schlegelii east of the Vogelkop 2-3. 

(19) Auriculars grey. Morotai to Batjan B6-7 ; Vanikoro G38 ; P. schlegelii 
east of the Vogelkop 2-3. 

(20) Mantle grey. Tasmania and Victoria F29~3oa (wholly grey) ; south- 
western and South Australia and New South Wales F28 & 30b (individuals with olive 
wash on rump or scapulars) ; midwestern Australia E22a-c (rump olive) ; north- 
western Australia E22d (rump and lower back olive). 

(21) Tail grey. Tasmania and southern Australia F28-30. 

(22) Mantle pure olive or grey (without phaeomelanins). Sumba and 
Salayer Ai & 5 ; Morotai to Batjan B6-7 ; San Cristobal C17 ; midwestern Australia 
to dahli and Snow Mountains E22-25 & 27 ; Vanikoro G38 ; Tabar H50 ; Tonga 
H57 ; P. schlegelii in Cyclops Mountains 2. 

ZOOL. 4, 4. II 



148 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

(23) Upperparts brown. Northern and central Fiji D18-21 ; Ceram H43, 
Louisiades H47, Mussau £L|.8d, Lihir H49. 

The mantle is distinctly brownish-olive on Aneitum and from Mai to Santo G36 
& 37c-d, from Timor to the Tenimber Isles H39-42 (tail pure olive on Babar H40), 
from Bum to the Peleng Isles H44-46, in the rest of the Bismarcks and on Manus 
H48a-c & 51, on Ndeni, Santa Cruz islets and Utupua H52-53, in southern Fiji 
H54-56, and in P. soror in the Vogelkop 1. 

(24) Upperparts sometimes rufous- washed. Choiseul to Guadalcanal, and 
Vella Lavella to Tetipari Cio-11 & 13-14 ; northern and central Fiji D 18-21. 

(25) Wings russet-edged. Solomons except San Cristobal C9-16 (mixed with 
olive from Choiseul and Kulambangra to Florida Cio & 15) ; northern and central 
Fiji D18-21 (mixed with olive on Koro D2ob). 

(26) Tail partly black. San Cristobal C17 ; north-western and northern 
Australia, dahli and Snow Mountains E22d-23, 25 & 27 ; P. soror. 

The variable hybrid populations on the small islands off Shortland E26 have 
largely been omitted from the character-geography. They are discussed on p. 156. 



NATURAL GROUPS 
Co-variation and character-complexes 

Single characters might be used to link populations into as many sets of overlapping 
assemblages as there are characters under consideration. If these bore no relation to 
one another, or were strictly correlated with environmental features, little could be 
inferred about relationships and evolutionary history. In many groups, the few 
characters available in ordinary museum material may be too sporadic or too liable 
to parallel evolution to be helpful. But where there are many characters, capable of 
independent variation and not exclusively related to environmental differences, 
major discontinuities can be detected despite the local elimination or independent 
origin of single characters. 

In the P. pectoralis superspecies, both the male and the female patterns are subject 
to a great deal of variation. The distribution of each character is different, so that 
all are at least partly independent. They are of very unequal systematic value ; 
great when their boundaries coincide with major discontinuities in the constellation 
of characters, small when they occur in forms not otherwise connected. Thus the 
loss of the male gorget is an important character, since in other respects also the 
Malaitan race C12 connects with those of northern Fiji D 18-19, and these with P. 
flavifrons. Hen-feathering, on the other hand, is unimportant, since the hen-feathered 
races A5, C16 and F33 resemble, not one another, but the females of neighbouring 
dimorphic forms. A character may be important in one part of the range though not 
in another. For example, grey-winged males distinguish the closely-related forms 
(group E) which range from midwestern Australia to the Bismarcks E22-25, from 
others whose ranges they approach closely and whose males are otherwise very similar 
(F31, H47-50, P. soror 3) : but such grey wings are found also in eastern Fiji D19 
& 21 & H56, in Tasmania and southern Australia F28-30, on Damar H41 and in P. 
soror 2 ; and not in the Snow Mountain race E27, which also belongs to group E. 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 149 

Striking variants of the male pattern, which seem to be relatively stable, mostly 
mark off local groups of populations with rather compact geographical boundaries. 
However, many of them are associated with very different character-complexes in 
different parts of the species range, and even locally a single character is seldom 
precisely co-extensive with the complex. The major discontinuities cannot be ade- 
quately defined by using these few more or less clear-cut characters alone. The much 
greater number of obviously quantitative differences must be considered as well. 
Although all are liable to parallel evolution and extensive intergradation, discon- 
tinuities are marked by concordant changes in a number of characters. 

Intergradation 

Few of the discontinuities are sharply defined. Almost everywhere, very distinct 
forms are connected by populations which are intermediate in range and character. 
Sometimes the changes are more or less regular in all the characters concerned, 
sometimes they are abrupt or out of step. At a few points (p. 160) the intermediate 
populations are highly variable, and it is clear that the intergradation is secondary — 
forms which differentiated in isolation have met and exchanged genes in a hybrid 
zone. Where the individual populations do not show exceptional variability, the 
intergradation may be primary or secondary. Primary intergradation could result 
from selection in relation to environmental gradients, or from incomplete isolation 
between diverging populations, or (conceivably) from the expression of orthogenetic 
trends after successive expansions of range (cf . Mayr & Moynihan, 1946, 1) . Secondary 
intergradation without increased variability would imply that the hybrid populations 
had been stabilized by subsequent selection. In P. pectoralis the geographical 
patterns of intergradation, and the characters involved, make it seem highly probable 
that the most striking examples of intergradation have resulted from the hybridizing 
of differentiated forms, with subsequent stabilization (p. 160). In other areas, how- 
ever, character gradients seem to be correlated with climatic differences (p. 186). 

Subspecies-groups 

There are all degrees of phenotypic discontinuity, from differences between 
individuals of the same population upwards. The precise scope and rank of any 
natural group of populations must be to some extent a matter of opinion, except that 
sympatry introduces an objective criterion at the species level. The useful scope of the 
subspecies is discussed on p. 175. For convenience, such a richly diverse species as 
P. pectoralis must be broken up into subspecies-groups. These will not be of equal 
distinctiveness and homogeneity. Where a rather compact range is occupied by 
closely related forms, yet divided by considerable discontinuities, the number of 
groups to be recognized is to some extent a matter of choice. For example, the 
Australian subspecies-groups E and F might be combined into one, since the 
differences between them are much less sharp than those which mark off some other 
groups (cf. Mayr, 1954a, 19). 

Where unlike forms intergrade they should not be separated if the connecting 
cline seems to be environmentally determined. Mathews (1930) not only removed the 



150 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

Northern Australian group E from P.pectoralis but split it into two species, on the basis 
of characters which strongly suggest selective adaptation to climate (p. 187). But 
where the intergradation seems to be secondary and the end-forms are sufficiently 
unlike, they should be separated even though the position of the dividing line will 
have to be arbitrarily decided. 

The situation in P. pectoralis is extremely complex. It seems to result from great 
plasticity in plumage characters combined with unusual ethological and genetic 
tolerance, and high mobility in successive waves of colonization combined with 
philopatry in local populations. As a result, very unlike forms have evolved and then 
met and interbred freely, and the resulting character-gradients mimic the true 
adaptive clines which also occur. It is unlikely that two students will ever agree on 
every detail of this confused situation. 

In the following descriptions, the superspecies has been divided into eleven groups. 
These are treated in an order determined by higher grouping according to characters 
and trends (p. 168), regardless of the implications of sympatry and intergradation. 
The use of binomens for some of these groups (e.g. " P. schlegelii ") and informal 
designations for others (e.g. " Solomons group C ") anticipates the findings of later 
sections. A description of the divergence of each group as a whole from the standard 
patterns is followed by character-geographies of departures from the group patterns 
so determined. In general, group patterns have been arrived at (like the standard 
patterns of the whole superspecies) by combining those characters which are found 
in a majority of the contained forms, excluding those which are obviously intermediate 
with other groups. But in the Lesser Sunda Isles, Moluccas and Fiji, it seems clear 
that few forms have escaped contamination by gene-exchange. Here the group 
patterns are taken to be those of the apparently pure forms Ai, B6-7, and Di8a & 19a. 

P. schlegelii (Text-fig. 3, p. 167) 

Small birds. Male chin black, throat-patch small ; gorget very broad, fore-edge 
defined by white feathers ; breast, flanks and belly rufous- washed ; mantle very 
dark ; wing, upper tail-coverts and tail black. Female throat barred grey and white, 
gorget grey, breast and flanks olive ; belly lemon-yellow ; cap and auriculars grey, 
mantle pure olive. 

Male Variation. Throat-patch larger and gorget narrower in Vogelkop 1. 
Rufous very deep in south-eastern New Guinea 3b, paling steadily westwards to 3a ; 
much paler in Vogelkop and Cyclops Mountains 1-2. 

Female variation. Throat white with grey bars, gorget and breast pale in 
Vogelkop and Cyclops Mountains 1-2 ; throat grey with white bars, gorget and 
breast dark from Weyland Mountains to south-eastern New Guinea 3. 

Greys and olives brownish in Vogelkop 1. 

Lesser Sundan group A (Text-fig. 2, p. 161) 

Very small birds. Male chin black, throat-patch small and separated by black 
feathers from auriculars ; gorget narrow, fore-edge defined by white feathers ; 
breast, belly and flanks rufous- washed ; collar narrow and olive-washed ; upper 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 151 

tail-coverts and tail black. Female throat whitish ; gorget greyish, pale and vague ; 
cap sandy-grey ; mantle pale sandy-olive. 

Male variation. Hen-feathered on Salayer A5 — male differs from female in 
larger throat-patch, greyer gorget and cap, grey auriculars, and darker and greener 
mantle. 

Throat-patch touches auriculars from Java to Alor and in Flores Sea A2-4 ; 
patch not small, and fore-edge of gorget with black tips, in Flores Sea A4. 

Rufous pale on belly in Java A2a, more so on Bali A2b, restricted to band behind 
gorget from Sumbawa to Alor A3, very pale and barely detectable below breast in 
Flores Sea A4. 

Wing black and mantle mottled with black in Flores Sea A4 (individually variable) . 

Tail with narrow pale tips from Java to Alor and in Flores Sea A2-4. 

Female variation. Throat pure white in Flores Sea and on Salayer A4-5, 
pinkish-buff from Java to Alor A2-3. 

Gorget rather pinkish from Java to Alor and on Salayer A2-3 & 5 ; breast and 
belly uniformly pinkish in Flores Sea A4. 

Breast and belly with very little yellow from Java to Alor and in Flores Sea A2-4. 

Cap almost pure grey and mantle pure olive on Salayer A5, distinctly sandy from 
Sumbawa to Alor and in Flores Sea A3-4, less so on Sumba and Java A1-2. 

Except for their smaller size, females from the Flores Sea A4 are very like those of 
Timor H39. 

Moluccan group B (Text-fig. 2) 

Medium-sized to large birds. Male chin black ; throat feathers long, partly 
covering gorget and cutting it off from auriculars ; gorget narrow from side to side, 
without black tips in the fore-edge ; upper tail-coverts and tail black. Female 
throat barred grey and white, gorget grey, breast and flanks pale olive ; belly lemon- 
yellow ; cap and auriculars grey, mantle pure olive. 

Size variation. Large birds on Ternate, Tidore and Obi B7-8. 

Male variation. Chin white, throat feathers shorter, and tail with narrow and 
obscure pale tips on Obi B8. 

Female Variation. Throat-bars and gorget buff, breast and flanks ochraceous, 
belly golden-yellow, cap and mantle brownish on Obi B8. 

Greys slightly paler and olives yellower on Ternate and Tidore B7 than from 
Morotai to Bat j an B6. 

The Obi form B8 is precisely intermediate between those of Ternate and Tidore 
B7 and Ceram H43, except that the female belly is deeper yellow than in either. 

Solomons group C (Text-fig. 2) 

Very large birds. Male chin black ; throat-patch yellow, rather small ; gorget 
broad, fore-edge defined by yellow feathers ; collar washed with olive. Female bill 
pale ; gorget broad and vague ; breast and flanks washed with melanins ; cap olive ; 
wing-feathers edged with russet. 

Size variation. Small birds on Rennell and San Cristobal C16-17, large (not very 
large) from Vella Lavella to Gatukai C13 & 15. 



152 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

Male variation. Hen-feathered on Rennell C16 — male almost indistinguishable 
from female, but with slightly more carotenoid on the average. 

Chin yellow on San Cristobal C17. 

Throat-feathers long (extending behind auriculars), gorget very broad on Vella 
Lavella and Ganonga C13. 

Gorget absent, but throat-feathers sometimes black-fringed, on Malaita C12. 

Some black tips in fore-edge of gorget on San Cristobal C17. 

Rufous patch below gorget on Santa Anna Ci7b and in individuals on San Cristobal 
Ci7a. 

Head olive with blackish lores and auriculars on Santa Anna Ci7b, varies from 
olive (auriculars brownish) to black on San Cristobal Ci7a. 

Collar absent on Vella Lavella and Ganonga C13 ; reduced to a trace on Malaita 
and from Kulambangra to Gatukai and Tetipari C12 & 14-15 ; reduced to a trace on 
the hind-neck, but with broad patches laterally, on San Cristobal C17. 

Entire upperparts and flanks black on Vella Lavella and Ganonga C13 (individuals 
on Ganonga Ci3b have narrow olive edges on the wings) ; wing black and mantle 
more or less mottled with black on Rendova and Tetipari C14 ; centres of mantle- 
feathers blackish, and olive dark greenish, from Kulambangra to Gatukai C15 ; 
olive rather greenish on San Cristobal C17. 

Upper tail-coverts olive from Kulambangra to Gatukai and on San Cristobal 
C15 & 17 ; olive edges broad on Malaita C12. Tail more or less olive on San Cristobal 

ci 7 . 

Female variation. Bill straw-coloured from Kulambangra to Guadalcanal and 
Rennell Cioc-11 & 15-16 ; blackish brown elsewhere. 

Individuals on Vella Lavella Ci3a are melanistic, with black gorgets and more or 
less black upperparts. 

Underparts faintly washed with yellow, cheeks yellower than throat, on Bougain- 
ville and Guadalcanal (juveniles sometimes much yellower) and Malaita Co,, 11 & 12 ; 
yellow pale on Rendova, Tetipari and Rennell C14 & 16 ; lemon-yellow elsewhere 
(when unmixed with melanins). 

Throat faintly barred on Malaita C12 ; underparts (except mid-belly) conspicuously 
shaft-streaked on Malaita C12, more faintly on Bougainville, Choiseul and Guadal- 
canal, and from Kulambangra to Gatukai Co/-ioa, n & 15. 

Gorget greyish on Bougainville and Malaita Co, & 12 ; olive from Choiseul and 
Kulambangra to Florida, and on Rennell and San Cristobal Cio & 15-17 (fairly 
narrow and distinct on San Cristobal C17) ; rufous on Guadalcanal and from Vella 
Lavella to Tetipari Cn & 13-14. 

Underparts washed with deep rufous on Rendova and Tetipari C14 ; rufous very 
variable in extent and depth on Vella Lavella and Ganonga C13 ; confined to gorget, 
breast and flanks on Ysabel and Guadalcanal Ciob & n (deep, pale or absent), and 
Choiseul, Russel Isles and Rennell Cioa & c & 16 (pale). 

Breast and flanks pale olive from Kulambangra to Gatukai and on Rennell 
C15-16, and in individuals from Choiseul to Florida Cio. 

Cap citrine, mantle olive on San Cristobal C17 ; cap brownish or rufous olive, 
mantle olive from Kulambangra to Gatukai and on Rennell C15-16 ; cap brownish 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 153 

citrine, mantle citrine from Buka to Malaita C9-10 & 12 (sometimes with a rufous wash 
on Ysabel Ciob, less common on Choiseul Cioa) ; cap rufous, scapulars and mantle 
strongly washed with rufous (most consistently on Rendova and Tetipari C14) on 
Guadalcanal and from Vella Lavella to Tetipari Cn & 13-14. Indications of a collar 
from Buka to Guadalcanal and on Vella Lavella and Ganonga C9-11 & 13. 

Olive dark from Kulambangra to Gatukai C15, darker still on Rennell C16. 

Olive dull (little carotenoid) on Rennell C16 ; richer on Guadalcanal, Rendova and 
Tetipari Cn & 14 ; richer on Bougainville and from Kulambangra to Gatukai Co, 
& 15 ; richer from Choiseul to Malaita and on Vella Lavella and Ganonga Cio, 12 
& 13 ; rich on San Cristobal C17. 

Wing-feathers edged with olive on San Cristobal C17 ; russet mixed with olive 
from Russel Isles to Kulambangra Cioc & 15, less strongly from Choiseul to Florida 
Cioa-b. 

Tail pure olive, with variable black subterminal patch, on San Cristobal C17 ; 
citrine from Buka to the central Solomons and Guadalcanal C9, 13-15 & ioc-ii ; 
olivaceous brown from Choiseul to Malaita Cioa-b & 12 ; dull brown on Rennell C16. 

Fijian group D (Text-fig. 2) 

Medium-sized birds. Male chin and throat yellow ; no gorget ; forehead yellow ; 
black of cap continued on to hind-neck, collar reduced to a trace ; olive very dark ; 
upper wing-coverts yellow-edged ; upper tail-coverts olive ; olive tips of tail-feathers 
dull but wide. Female forehead, circumoculars, cheeks and under tail-coverts 
faintly washed with yellow, underparts otherwise without carotenoid (buffy-grey, 
with throat and mid-belly slightly paler) ; underparts streaked with brown, and throat 
and breast barred also ; upperparts dull olive-brown, wing-feathers edged with dark 
russet ; individuals are dark rufous all over. 

Male variation. Underparts lemon-yellow on Viti Levu and Ovalau D18, 
orange-yellow elsewhere. 

More or less complete gorget, individually variable, on Ovalau and from south- 
western Vanua Levu to Kio Di8b & igb-c ; gorget complete (though somewhat 
irregular) on Taviuni, Koro and Vatu vara D20-21. 

Breast, belly and under tail-coverts washed with brown on Vatu vara D21 ; under 
tail-coverts ochraceous on Koro D2ob, decreasingly so towards Vanua Levu D2oa to 19b. 

Collar narrow but uninterrupted on Vatu vara D21. 

Wing-quills edged with greyish on Vanua Levu and Vatu vara D19 & 21, becoming 
more olive eastwards, towards Viti Levu and Koro D18 & 20. 

Upper tail-coverts with black centres on Koro and (larger) on Vatu vara D20D-21. 

Female variation. Bill brown on Rambi and Kio D19C. 

Underparts evenly-coloured on Koro and Vatu vara D2ob & 21 ; mottling becomes 
more common and emphatic towards the large islands Di8b to a, and 20a to 19a. 
Ground-colour of underparts cinnamon on Koro D2ob and (rather paler) Vatu 
vara D21, individually variable, becoming greyer, towards the large islands Di8b 
to a, and 20a to 19a. 

Forehead, circumoculars and cheeks not yellow on Koro and Vatu vara D20D-21 ; 
increasingly so towards the large islands Di8b to a, and 20a to 19a. 



154 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

Females on Koro and Vatu vara D20D-21 are very like those in southern Fiji 
H54-56. 

P.flavifrons (Text-fig. 5, p. 173) 

Small birds. Male chin dull black, throat dull black with broad j^ellow or white tips; 
no gorget ; underparts lemon yellow ; forehead yellow or white ; upperparts dull 
black (slightly washed with olive), primaries with obscure narrow greyish edges. 
Female cock-feathered — like the male, but throat pale grey with narrow yellow or 
white tips, forehead blackish (with or without a yellow wash) . 

Individual variation. The tips of the throat feathers are usually yellow, but 
may be white ; two specimens in the British Museum (Natural History) have mixed 
yellow and white tips. Some white-throated birds have the forehead white (in 
females, without a yellow wash) . The distribution of these phases in the collections 
of the British Museum (Natural History), and the American Museum of Natural 
History (from Mayr, 19326), is shown in Table II : there is no evidence of a significant 
difference in representation of the colour phases, either between the sexes or between 
Savaii and Upolu. All the B.M. (N.H.) specimens are from Upolu. 

Table II. — Colour Phases in P. flavifrons. 

Forehead. 

A 





Throat. 




r 

Yellow. 


Yellow 


. . . 


. 


48 


Yellow and white . 


. 


2 


White 




. 


8 



White. 



P. soror (Text-fig. 3) 

Small birds. Male throat-patch rather small ; gorget narrow ; underparts lemon- 
yellow ; breast and flanks faintly olivaceous ; collar reduced to a trace ; mantle 
very dark greenish olive ; edges of wing quills greyish ; upper tail-coverts olive, 
tail partly olive. Female throat-patch white and small ; gorget narrow, clear and 
brownish ; breast and flanks pale citrine, yellow wash extending high on breast ; 
belly pale lemon-yellow ; cap olivaceous-brown ; tail partly black. 

Male variation. Mantle increasingly golden-olive towards the west 3 to 1. 
Wing quills greyest in the centre 2. 

Tail olive, with a variable black subterminal patch, in the west and east 1 & 3 ; 
black, with wide olive edges basally, in the centre 2. 

Female variation. Upperparts much browner in Vogelkop 1 ; increasingly 
purer olive towards the east 2 to 3. 

Northern Australian group E 

Very small to medium-sized birds. Male throat-feathers white to their bases ; 
edges of wing-quills grey ; edges of upper wing-coverts yellow ; primary-coverts 
black. Female throat white, with barring ; gorget well defined, buffy-grey ; breast, 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 155 

flanks and belly whitish to golden-yellow, without melanins ; cap rather pure 
grey, mantle pure olive (or grey) ; tail partly black. 

Size variation. Very small birds in mid-western Australia E22a-c, size increasing 
through north-western Australia E22d to small in northern Australia and southern 
New Guinea E23-24. Like the Snow Mountain birds E27, those grouped together as 
dahli E25 are mostly medium-sized — but there is considerable variation between 
the scattered populations, with the birds of Fergusson E25C (and Teste and Long 
Islands E25b & d) rather large, and those of south-eastern New Guinea and Witu 
E25a & e apparently rather small. 

Male variation. Throat-feathers grey-based in Snow Mountains E27 (Text-fig. 3) . 

Breast and belly lemon-yellow in mid-western Australia E22a, gradually becoming 
deeper northwards E22b to d, golden-yellow from northern Australia eastwards E23-27. 

Collar narrow in midwestern Australia E22a (widening northwards E22bto23), 
and Snow Mountains E27. 

Mantle greenish-olive in midwestern Australia E22a (becoming yellower north- 
wards E22bto23), and Snow Mountains E27. 

Edges of wing-quills olive in Snow Mountains E27. 

Primary-coverts olive in mid-western and north-western Australia and Snow 
Mountains E22 & 27 ; sometimes with very narrow pale edges from northern Australia 
to southern New Guinea E23 to 24. 

Upper tail-coverts olive in mid-western and north-western Australia E22. 

Tail with broad olive edges basally in midwestern Australia E22a, decreasing 
northwards E22b to d. 

Female variation. Females from Teste and Fergusson E25D-C are unknown. 

Throat unbarred in mid-western Australia E22a-c, barring pale in north-western 
and northern Australia E22d-23 ; throat and gorget shaft-streaked (in addition to 
barring) from Cape York to southern New Guinea E24 (faintly) and in dahli and 
Snow Mountains E25 & 27. 

Breast and flanks pale olivaceous in Snow Mountains E27 ; ochraceous from Cape 
York to southern New Guinea E24. 

Breast and belly whitish and under tail-coverts pale yellow in mid- western Australia 
E22a-c ; breast and belly pale yellow, under tail-coverts lemon-yellow, in north- 
western Australia, and from Cape York to southern New Guinea E22d & 24 ; under- 
pays lemon-yellow in northern Australia E23, golden-yellow in dahli and Snow 
Mountains E25 & 27. 

Cap brown in Snow Mountains E27. 

Mantle grey, only rump olivaceous, in mid-western Australia E22a-c, becoming 
olive northwards E22dto23. Olive bright in Snow Mountains E27. 

Edges of wing feathers, rump and base of tail brownish on Witu, Malie and Nissan 
E25e & j-k. 

Tail olive in mid-western Australia, and from Cape York to southern New Guinea 
E22a-c & 24 ; with a variable amount of black in north-western and northern 
Australia E22d-23 ; more than half black in dahli E25 ; black with olive edges in 
Snow Mountains E27. 

Mayr has shown (1932a) that the birds on small islands west of Shortland E26 



156 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

form a hybrid population. The nine known males show almost every degree of inter- 
mediacy between dahli E25 and the Bougainville-Shortland race Co, — except that 
they do not have black chins like those of the Solomons group C. In this series the 
largest birds have the yellowest throats, and the smallest ones the whitest. Other 
characters seem to vary independently, so far as can be judged from this short 
series. There is rather less variation between the nine females, which are also inter- 
mediate between those of E25 and C9. 

The females from Cape York to southern New Guinea E24 differ from those on 
either side (E23 & 25) in characters which approach those of the Queensland form 
F31 — breast and flanks ochraceous, little ventral yellow, tail without black (Mayr, 
1954a). 

The Louisiades form H47 is intermediate between groups E and H. It is here 
placed in the latter because the male has olive edges to the wings, and the female is 
brownish ; but the white throat, sharp and narrow gorget and bright yellow under- 
pays of the female are untypical of group H. The Tabar form H50 may be similarly 
intermediate (p. 164). 

Southern Australian group F 

Large birds, with long wings and tails, small bills and soft plumage. Male breast 
and belly lemon-yellow ; collar rather narrow and washed with olive ; upper wing- 
coverts yellow-edged ; upper tail-coverts olive, base of tail olive or grey. Female 
throat barred ; gorget, breast and flanks buffy-grey, belly more or less washed with 
melanins ; yellow absent from underparts, or only in under tail-coverts ; cap grey, 
mantle dull sandy-olive. 

Size variation. Large birds with long tails and very small bills in Tasmania 
F29, size and relative tail-length decreasing and bills lengthening northwards — 
smallest birds near Cairns F3ib, longest bills on Lord Howe and Norfolk Islands 

F32-33. 

Male variation. Hen-feathered on Norfolk Island F33 — indistinguishable from 
female. 

Yellow somewhat paler from south-western Australia to Tasmania F28-29. 

Collar broader and clearer on Lord Howe F32. 

Wing quills blackish brown with grey edges in Tasmania F29, centres blackening 
and edges becoming more olive northwards F28 and 30 to 31. 

Tail without black in Tasmania F29 ; about one-third black from south-western 
Australia to South Australia F28a-b ; increasing through Victorian mallee F28C 
to two-thirds black in south-eastern Australia F30 ; two-thirds black on Lord Howe 
F32 (olive tips very wide) ; mainly black, olive varying from wide edges to a wash 
at the extreme base, in Queensland F31 (averaging blacker near Cairns F3ib). 

Pale part of tail grey in Tasmania and southern Australia F28-3oa ; olive else- 
where, but often greyish near Cairns F3ib. 

Female variation. Belly pinkish from south-western Australia to South 
Australia F28a-b, fading to buffy in Victorian mallee, Tasmania and near Cairns 
F28C-29 & S 1 ^ i whitish in south-eastern Australia F30 ; often faintly yellowish in 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 157 

southern Queensland F3ia, and always on Lord Howe F32 ; distinctly yellow on 
Norfolk Island F33. 

Under tail-coverts whitish in Tasmania and southern Australia F28-30. 

Upperparts rather brownish in Tasmania F29, and more or less brownish in most 
individuals near Cairns F3ib ; rather pure grey in south-western Australia F28a. 

Mantle grey in Tasmania F29 ; sometimes with a faint olive wash in southern 
Australia F28 & 30 ; dull olive elsewhere. 

Tail grey in Tasmania and southern Australia F28-30 ; dull olive near Cairns 
F3ib ; olive elsewhere. 

Southern Melanesian group G 

Small birds. Male gorget narrow ; breast and belly lemon-yellow ; collar narrow 
and washed with olive ; centres of wing-feathers blackish-brown ; upper tail-coverts 
and tail olive. Female throat white ; gorget greyish ; breast and belly without 
melanins. 

Size variation. Very large birds on Loyalty Islands G35. 

Male variation. Lateral throat feathers long, separating gorget from auriculars, 
on New Caledonia G34 (Text-fig. 5). 

Gorget dull black, and very narrow, on New Caledonia G34 ; dull black on Aneitum 
G36 ; rather broad on Vanikoro G38. 

Pale rufous patch below gorget on New Caledonia G34. 

Flanks and under tail-coverts ochraceous, rump, tail and edges of secondaries 
washed with brownish, in individuals on Aneitum G36. 

Breast and belly golden-yellow on Loyalty Islands G35. 

Cap and auriculars grey on New Caledonia G34 ; dull black, cap more or less 
scalloped with olive, on Aneitum G36. 

Collar reduced to a trace on New Caledonia G34, very narrow on Erromango G37a. 

Mantle greenish-olive on Erromango, and from Raga to Vanikoro G37a & 376-38 ; 
golden-olive on Loyalty Islands, and Malekula and Santo G35 & 3yd. 

Upper tail-coverts black-centred, tail black, on Vanikoro G38 ; tail with black 
patches (on inner webs of all but central quills) from Erromango to Banks Islands 

G37. 

Female variation. Throat and gorget shaft-streaked on Vanikoro G38. 

Gorget olive on Loyalty Islands and Vanikoro G35 & 38 ; greyest on Erromango, 
Efate and Banks Islands G37a-b & f ; brownish on Aneitum G36. 

Gorget extends into buffy-grey wash on breast and belly on New Caledonia G34. 

Underparts golden-yellow on Loyalty Islands G35, lemon-yellow on Vanikoro G38 ; 
elsewhere, under tail-coverts pale yellow, breast and belly faintly washed with yellow 
(palest on Malekula and Santo G37d). 

Cap and auriculars grey on Vanikoro G38 ; cap greyish on Efate and Banks 
Islands G37b & f ; brownish on Aneitum G36. 

Mantle pure golden-olive on Loyalty Islands G35, pure greenish-olive on Vanikoro 
G38 ; elsewhere dull olive, purest on Efate and Banks Islands G37b & f , brownish on 
Malekula and Santo G37d ; very brown on Aneitum G36. 



158 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

Tail and edges of secondaries browner than mantle on Loyalty Islands G35, some- 
what so on Aneitum and Erromango G36-37a. 

Juvenile variation. Throat lemon-yellow on Vanikoro G38, sometimes yellow- 
washed in New Hebrides (recorded from G37C & f by Mayr, 19326). Juveniles on 
Vanikoro G38 are remarkably like those on San Cristobal C17, except for the shaft- 
streaking of their throats and gorgets. 

The New Hebrides G36-37 are a region of incipient subspeciation, with the females 
of almost every island distinguishable in series (Mayr, 19326). The Aneitum popula- 
tion G36 is rather distinct, and that on Erromango G37a more so than the others. 

Widespread group H (close to standard patterns, Text-fig. 1) 

Small to very large birds. Male of standard pattern. Female underparts rather 
uniform, brownish, with little yellow ; upperparts brownish, cap not contrasting, 
edges of wing feathers distinctly browner than mantle. 

Size variation. Very large birds on Tenimber Isles, Lihir and Tonga H42, 49 
& 57 ; large on Tabar and Lau archipelago H50 & 56 ; small on Timor and Damar, 
Sula and Peleng Isles, large islands in the Bismarcks, and Kandavu H39, 41, 45-46, 
48 & 54. 

Male variation. Chin and throat black in Tonga H57 (Text-fig. 5) ; yellow on 
Tenimber Isles H42. 

Throat-feathers long, cutting gorget off from auriculars (fore-edge of gorget defined 
by white feathers) on Ceram, and Ndeni and Santa Cruz islets H43 & 52 ; gorget 
very narrow near auriculars, with few black tips, on Buru H44. 

Gorget streaked with white on Babar H40 ; dull black and very narrow on Kandavu 
H54, narrow on Ngau H55. 

Underparts lemon-yellow from Timor to Tenimber Isles and on Utupua, Kandavu 
and Tonga H39-42, 53-54 & 57 ; orange-yellow in Louisiades and on Lihir and Manus 
H47, 49 & 51. 

Under tail-coverts ochraceous on Ngau H55. 

Upperparts (except for narrow olive edges on wing-feathers) black on Ndeni and 
Santa Cruz islets H52 ; mantle mottled with black in Lau archipelago H56, somewhat 
so on Ngau H55. 

Primary-coverts black on Ceram and Buru, Ndeni and Santa Cruz islets, and Lau 
archipelago H43-44, 52 & 56. 

Collar absent on Ndeni and Santa Cruz islets H52 ; reduced to a trace on Tenimber 
Isles H42, and Ngau and Lau archipelago H55-56 (black of cap extends on to hind- 
neck) ; washed with olive from Timor to Damar, in the Louisiades and on Utupua 
and Kandavu H39-41, 47 & 53-54. 

Olive dark and greenish on Buru, Ngau and Lau archipelago H44 & 55-56 ; dull 
on Kandavu H54. 

Olive washed with brownish (especially on rump, tail and edges of secondaries) 
on Ceram and Louisiades H43 & 47. 

Edges of wing quills grey on Damar H41, and greyish in Lau archipelago H56. 

Upper tail-coverts olive on Timor, Tenimber Isles, Louisiades, Kandavu and Tonga 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 159 

H39, 42, 47, 54 & 57 ; black from Ceram to Peleng Isles H43-46 (very narrow olive 
edges on Buru H44). 

Tail olive on Timor and western Louisiades H39 & 47a ; olive with blackish 
subterminal patches on Misima and Kandavu 1147b & 54 ; olive basally on Babar 
and Rossel H40 & 47c ; with broad olive edges basally on Damar H41. 

Pale tip of tail very broad and yellowish in Tonga H57. 

Female variation. Throat white on Timor and Babar, Louisiades and Utupua 
H39-40, 47 & 53 ; whitish on Damar, Ceram, Sula and Peleng Isles, Tabar, Manus, 
Ndeni and Santa Cruz islets H41, 43, 45-46 & 50-52. 

Throat washed with yellow on Tenimber Isles H42. 

Throat barred on Damar, Sula and Peleng Isles H41 & 45-46, and faintly in Tonga 

H57- 

Throat and gorget streaked on Manus H51, and faintly on Tabar H50. 

Underparts uniformly cinnamon in southern Fiji H54-56 (very richly so on Ngau 
H55). Gorget scarcely darker than throat and belly on Mussau and Lihir H48d~49 ; 
little darker on Tenimber Isles, large islands of the Bismarcks, and Ndeni and Santa 
Cruz islets H42, 48a-c & 52. Gorget little darker than breast, but greyer, on Timor 
and Damar and from Ceram to Peleng Isles H39, 41 & 43-46. Gorget pale and 
narrow on Babar and Louisiades H40 & 47 (vinous), Tabar, Manus and Tonga 
H50-51 & 57 (cinnamon) ; broad on Utupua H53 (dark brown). 

Breast and belly without melanins in Louisiades H47 ; belly without melanins, 
breast faintly cinnamon on Tabar and Tonga H50 & 57 ; belly without melanins, 
breast and flanks brownish on Utupua H53 ; breast and belly faintly washed with 
cinnamon on Babar and Manus H40 & 51. 

Breast and belly without yellow on Ngau and Lau archipelago H55-56 ; faintly 
washed with yellow on Timor, Damar, Buru and Kandavu H39, 41, 44 & 54, more 
strongly on Tenimber Isles, Ceram and Sula Isles H42-43 & 45 ; yellow pale on Babar, 
Bismarcks except Tabar, and Ndeni and Santa Cruz islets H40, 48-49 & 52 ; deep 
in Peleng Isles and Louisiades H46 & 47, and especially on Tabar and Manus 
H50-51. 

Cap grey on Tabar, Ngau and Tonga H50, 55 & 57 ; distinctly greyer than mantle 
on Manus, Utupua, Kandavu and Lau archipelago H51, 53, 54 & 56 ; distinctly 
browner than mantle on Bismarcks except Tabar H48-49. 

Mantle bright citrine on Tabar, Manus and Tonga H50-51 & 57 ; dull sandy- 
olive on Timor, Damar, Tenimber Isles, Kandavu and Lau archipelago H39, 41-42, 
54 & 56 ; brownish olive on Babar H40 (tail pure olive), Peleng Isles, Louisiades, 
large islands of the Bismarcks, Utupua and Ngau H46, 47, 48a-c, 53 & 55 ; browner 
and duller from Ceram to Sula Isles and on Ndeni and Santa Cruz islets H43-45 & 
52 ; olive-brown on Mussau and Lihir H48d~49. 

Edges of wing feathers scarcely browner than mantle on Tabar, Manus and Tonga 
H50-51 & 57. 

Not only are the distinctive characters of the group few and slight, but most of the 
forms are intermediate in some respects with neighbouring groups. Besides penetra- 
tion by characters proper to other groups, intermediate forms seem to be liable to 
special changes (p. 165). The wide range of this group has enabled it to meet and 



i6o THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

intergrade with five of the seven other subspecies-groups, and there seems to have 
been gene-flow into it in the following areas : 

From group A to Timor and Babar H39-40. 

From group B to Ceram, Peleng and Sula Isles H43 & 46-45. 

From group D to southern Fiji H54-56. 

From group E to Louisiades H47 (and possibly Damar, Tabar and Manus H41 

& 5o-5i). 

From group G to Utupua H53. 

When the effects of this gene-exchange have been allowed for, three forms stand out 
conspicuously from the remarkably uniform remainder : 

Tenimber Isles H42 (very large, throat yellow). 

Ndeni and Santa Cruz islets H52 (male upperparts black, throat-feathers long) . 

Tonga H57 (very large, male throat black, female with little brown and much 
yellow). 

INTERGRADATION AND BRIDGELESS GAPS 
Gene-exchange between subspecies-groups 

The eight subspecies-groups of P. pectoralis approach one another in several areas, 
in all of which there are signs of intergradation between them (Text-fig. 8) . Groups 
A and H intergrade in the Lesser Sunda Isles, B and H in the southern Moluccas, 
C and E in the western Solomons, C and G in the eastern Solomons, D and H in 
central Fiji, E and F in southern New Guinea, E and H in the Louisiades, and G and 
H in the Santa Cruz archipelago. In most of these areas, it is not immediately 
obvious whether the intergradation is primary or secondary. 

Only in the western Solomons is there indisputable evidence of hybridization 
between very dissimilar forms. The populations on three islets west of Shortland 
(E26) are highly variable. Extreme individuals closely resemble the Shortland race 
Co, on the one hand and dahli E25 on the other, while every degree of intermediacy is 
found (p. 156). 

On San Cristobal Ci7a also there is considerable variation between males, in the 
amount of rufous on the breast and of black on the head and tail. Their song, too, 
is remarkably variable (Cain & Galbraith, 1956). Though the San Cristobal race 
agrees with the rest of group C in the yellow throat, and the olive cap and pale bill of 
the female, it differs from the standard pattern of that group in most other characters. 
It is much smaller. The male has a yellow chin, black tips in the fore-edge of the 
gorget, a rufous breast-patch, and partly olive head and tail. The female is bright 
yellow underneath, with a narrow gorget and no melanic wash, lacks rufous on the 
upperparts (including the wings) , and has a partly black tail. Many of these characters 
of the males are retarded, and of the females advanced. 

Avifaunally San Cristobal is distinct from the rest of the Solomons, with several 
endemic forms showing markedly reduced size (Collocalia esculenta makirensis, 
Ptilinopus solomonensis solomonensis , Halcyon chloris solomonis, Rhipidura rufifrons 
russata, Monarcha vidua, Myiagra cervinicauda and Aplonis grandis dichrous), 
reduced sexual dimorphism (Coracina tenuirostris salamonis, C. lineata makirae, 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 



161 







Fig. 2. — Four extreme forms of P. pectoralis (schlegelii assemblage) which intergrade with 
more nearly standard ones (soror assemblage). Sumba Ai ; Ternate B7 ; Shortland 
C9 ; Vanua Levu Diga. 



1 62 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

Myiagra cervinicauda, Myzomela nigrita tristrami and Dicaeum tristrami), or increased 
variability (Ptilinopus solomonensis solomonensis , Halcyon chloris solomonis, Coracina 
lineata makirae and Petroica multicolor polymorpha) in comparison with their 
western representatives. These trends may result simply from the isolated peripheral 
position and impoverished avifauna of the island. However, several birds on San 
Cristobal and its outlying islands are derived from southern Melanesia rather than 
from the rest of the Solomons [Ptilinopus richardsii, Lalage leucopygia, Vitia parens, 
Rhipidura fuliginosa, Myiagra cervinicauda and Myzomela cardinalis). The vari- 
ability of the male of P. pectoralis here suggests the possibility of hybridization, which 
might have disturbed the sexual dimorphism (p. 165). The Vanikoro race G38 shows 
some of the characters to be expected in the other putative parent : the female is 
rather similar to that on San Cristobal, except for its streaked white throat, while 
the juvenile has a streaked but yellow throat. The San Cristobal race shares with 
most members of group G the pale tail of the male and the melanin-free throat of 
the female ; and with the New Caledonian race G34 the rufous breast-patch and pale 
cap. In the light of extensive hybridization between unlike forms elsewhere, it 
seems most probable that the San Cristobal race is a hybrid between groups C and 
G. 

The New Caledonian race G34 differs from its neighbours of group G in several 
systematically important characters of the male plumage, which suggest affinities with 
P. schlegelii and groups A to C of P. pectoralis (the schlegelii assemblage, p. 168). The 
elongation of the white feathers at the sides of the throat and the failure of the gorget 
to join the auriculars are reminiscent of group B. The absence of black tips in the 
fore-edge of the gorget (laterally) is characteristic of the whole assemblage. The 
rufous breast is found elsewhere only in P. schlegelii and group A, and on San Cristobal, 
and is in the form of a patch rather than a band only on New Caledonia and San 
Cristobal. These two forms share also the pale cap, and the suppression of the collar 
(except for lateral patches in the latter). Though the females on these two islands 
are very different, that on New Caledonia agrees with typical females of group C 
rather than of group G in having the belly washed with melanins. The number of 
characters involved makes it improbable that they have been developed independ- 
ently, and it seems likely that this race too has had genetic contributions from group C 
as well as group G. 

Two further forms, here assigned to group H, show characters which are elsewhere 
systematically important and confined to the schlegelii assemblage (see Table III). 
The race on the Tenimber Isles H42 has a yellow throat. That on Ndeni and the Santa 
Cruz islets H52 has overlapping throat feathers without black tips, and a black 
mantle. But in other respects these forms agree well with members of group H. It 
would not be justifiable (without further evidence) to deny the possibility of inden- 
pendent origin for these characters, and to postulate mixed ancestry for these forms 
also. A latent tendency towards yellowing of the throat is apparent in many forms 
(p. 142). In the melanic race H52 the elongation of the throat feathers and the 
exclusion of melanin from their tips may be significant in maintaining the con- 
spicuousness of the throat-patch despite encroachment by black areas (as in the 
other melanic race C13). 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 163 

The populations between Koro and the large Fijian islands (Di8b & io,b-2oa) 
are evidently hybrid (Mayr, 1932b). There is much variability — especially on Ovalau, 
Rambi and Kio Di8b & 19c — in the degree of development of the male gorget and 
forehead-spots, and in the colour and streakiness of the female underparts. The 
average character of the populations changes progressively from Koro to the main- 
land of Vanua Levu (D2ob to 19a) , while the population on Ovalau Di8b is intermediate 
in character between those of Koro D2ob and Vitu Levu Di8a. Unfortunately, my 
material is inadequate to demonstrate quantitatively the changes in hybrid index 
and in variability which are qualitatively apparent in Mayr's description. 

The races on Koro and Vatu vara D20D-21 are themselves intermediate in character 
between the extreme Fijian forms (Di8a & 19a) and members of group H, to which 
they are linked by the races in southern Fiji H54-56. Males on Koro and Vatu vara 
have yellow throats with gorgets, those in southern Fiji white throats with gorgets ; 
the yellow forehead reappears on Vatu vara ; while all Fijian males have the collar 
obscure or very narrow. Females in southern and central Fiji (D20-21 & H54-56) 
are all much alike — differing from those of northern Fiji in having unstreaked 
cinnamon underparts, and from most of those in group H in having them almost uni- 
form from chin to vent and with scarcely any carotenoid. Blackness of the mantle 
(D20 & H55-56) and greyness of the wing quills (D19 & 21 & H56) are common to 
forms on either side of the intergroup boundary. 

Group H intergrades with groups A and B also. The male on Sumba Ai has a 
black chin, a small throat-patch, no black tips in the fore-edge of the gorget, a rufous 
ventral wash and a wholly black tail. These characters are reduced or absent elsewhere 
in group A, the dilution being greatest in the Flores Sea A4 and least in Java A2a. 
Although there is a sharp change in size at the Ombai Strait (along which the boundary 
between groups A and H has been drawn), and the special male characters of group 
A do not appear east of it, the female on Timor H39 is very like that in the Flores 
Sea (which differs from other females of group A in having a pure white throat 
and pinkish belly), and there is a fairly complete series of forms leading to a typical 
member of group H on Buru (H39 to 41 & 44). 

Males in the northern Moluccas B6-7 have black chins, long throat-feathers 
partly covering the gorgets, and wholly black tails ; while the females are unusual 
in their heavily-barred throats, pure greys and olives, olive breasts and grey auriculars. 
There is a perfect series of intermediates leading from these through Obi B8 and 
Ceram H43 to Buru H44. Although the male characters of group B do not appear 
there, the female in the Peleng Isles H46 is very like that on Obi B8, and the Sula 
Isles female H45 is intermediate between those of Peleng and Buru. 

The changes involved are quite different in these three areas of intergradation. 
They involve oddities of pattern, as well as the quantitative changes which seem 
more likely to be subject to environmental selection. Only in the Lesser Sunda 
Isles is there a marked climatic gradient (of increasing aridity eastwards from Java 
to Timor) with which the progressive change in character might be correlated. But 
here the character-progression (from Ai-2a-2b-3-4-H39) does not run parallel to 
that of climate. In the Moluccas the changes span only seven degrees of latitude 
across the equator, and in Fiji four degrees within the tropics, so that regular climatic 

ZOOL. 4, 4. 12 



164 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

changes cannot be great. It seems unlikely that adaptation to environmental 
gradients is involved in these character-progressions, although adaptive clines 
might be developed in relation to subtle differences (perhaps in the fauna or flora). 
The geographical patterns of character-dilution suggest introgression rather than 
continued gene-exchange between gradually diverging populations (see Text-fig. 8). 
Finally, groups A, B and D seem (p. 168) to be more closely related to one another 
than to group H, with which they all intergrade. 

There is thus no reason to believe that the intergradation in these areas is primary. 
The situation can be explained with the maximum economy of hypothesis in terms 
of a recent burst of range expansion by group H, which has come into secondary 
contact with diverse and anciently-isolated forms and freely interbred with them. 
The same thing has almost certainly happened in the western Solomons, where 
dahli E25 is a relatively very recent arrival and the hybrid population E26 is still 
exceedingly variable. Meise (1936) and Mayr (1942) have suggested that the Pachy- 
cephala on Koro D2ob is of hybrid origin and has been genetically stabilized by 
subsequent selection. This seems much the most probable explanation for the other 
intermediate, though not especially variable, forms just considered. 

Since the characters which distinguish groups E to H from one another are 
relatively slight and mainly quantitative, it is more difficult to decide whether the 
intergradation between these groups is primary or secondary. On the other hand 
there is less reason to doubt that the rather similar forms involved can interbreed, 
and the character-geography strongly suggests gene-flow between these groups in 
several areas (Text-fig. 8). 

Mayr (1954a) has pointed out that, while the male in southern New Guinea E24 
is like its relatives E23 and 25, the female shows (in its buffy belly with little yellow, 
and lack of black in the tail) dilution by characters of group F. The dilution appears 
to decrease into south-eastern New Guinea E26a (Rand, 1940). It is clear that 
gene-flow from northern Queensland F3ib is involved. Mayr also suggests that the 
character gradient in mid-western Australia (E22d to a) is the result of gene-flow 
from south-western Australia F28a. The partly olive tail of the male and the reduc- 
tion of carotenoid support this ; but the birds become smaller southwards instead 
of larger, and the female shows no sign of the pinkish underparts and barred throat 
characteristic of the south-western form. It seems more probable that the changes 
are clinal (p. 187). 

In the Louisiades H47, the female shows the bright yellow underparts, white 
throat (though without mottling) and narrow gorget of group E, but has a very 
brownish mantle like those of group H, while the male has olive edges to the wing 
quills. This form could equally well be assigned to either of these groups. The 
female on Tabar H50 resembles those of group E in the same respects, as to a lesser 
degree does that on Manus H51. It seems reasonable to suppose that the inter- 
gradation in the Louisiades is secondary, and this might be true of the Tabar race 
also ; but it is most unlikely that the small and recently-arrived populations of 
dahli E25 in the Bismarcks could sufficiently swamp the presumably large population 
on isolated Manus. This is surely a case of convergence, as in Tonga H57. 

While the male on Damar H41 is very like those of Timor and Babar H39-40 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 165 

(which I suppose to have been affected by gene-flow from group A), the female is 
different, closely resembling that of the Sula Isles H45 (which has affinities with 
group B). There may perhaps have been gene-flow across the Banda Sea. But the 
barred throat characterizing these females is also found in Australia, and the male on 
Damar has grey edges to the wing quills like those of group E. If gene-flow (other 
than from group A) is involved here, it seems more likely that it has been across the 
Timor Sea from northern Australia E23. 

The three forms in the Santa Cruz archipelago are remarkably different. The male 
on Ndeni and the islets H52 is melanic and has the throat feathers elongated, while 
those on Utupua H53 and Vanikoro G38 are close to the standard. Groups E to H 
differ mainly in the characters of the female. The Ndeni female is very near the 
standard, and a typical member of group H ; that on Vanikoro is quite different, 
most like the Snow Mountain female E27, and apparently a northern representative 
of group G ; while that on Utupua is intermediate between them. It is possible 
that this intergradation is primary, and that group H has spread west and east 
after its origin in the Santa Cruz, but more probably the contact was secondary. 

Marks of hybridity 

Knowing that hybridization is possible between very dissimilar forms of P. pectoralis, 
we have considered many forms which are intermediate, both geographically and 
phenotypically, between well-marked groups to be of hybrid origin, even though their 
variability is not exceptional. Several of these presumably hybrid forms differ from 
both putative parents (and from most other forms in the superspecies) by characters 
which are retarded in the male, or advanced in the female plumage. Partly olive 
(or grey) tails in the male are found in P. soror and the southern members of groups E, 
F and G ; in certain presumptive hybrids (San Cristobal C17, Timor to Damar 
H39-41, Louisiades H47 and Kandavu H54) ; and nowhere else. Female throats 
without melanic washes or mottling are found in P. soror and the southern members 
of groups E and G ; in some hybrids (Flores Sea and Salayer A4-5, San Cristobal 
C17, Timor and Babar H39-40, Louisiades H47 and Utupua H53) ; and nowhere 
else. Partly black tails in the female are found in P. soror and members of group E ; 
on San Cristobal C17 (and black shafts on Babar H40) ; and nowhere else. The 
presumptive hybrid females on Obi B8, Babar H40 and Utupua H53 have much 
brighter yellow bellies than their relatives on either side. 

In other parts of the species range, the amount of black in the tails of both sexes, 
and the depth of carotenoid coloration, are involved in character-progressions 
which may be climatically determined (p. 186). But while in these clines intensity 
of pigmentation increases with increasing temperature and humidity, in the forms 
under discussion the male tail tends to be pale and the female tail and belly to be 
deeply-coloured. Furthermore, there is no reason to suppose that the islands in 
the Flores Sea, Timor, Babar, Damar, Obi, the Louisiades, San Cristobal, Utupua 
and Kandavu share environmental factors which distinguish them from neighbouring 
islands. If the apparent regularities are real, it seems most probable that they are 
related to the secondary intergradation which characterizes P. pectoralis in these areas. 

These trends are not the phenotypic expression of heterosis, since the variable 



i66 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

hybrid populations E26, Di8b & igb-c do not show them. Possibly the tendency 
towards breakdown of the sexual dimorphism reflects genetic disturbances produced 
by hybridization, or selective changes involved in regaining a balanced genome. 
These might be expected sometimes to blur the distinction between the adult female 
and juvenile plumages, producing retarded rather than advanced characters in the 
female. This may be the explanation for the reversal of south-to-north changes in 
northern Queensland (p. 156). Many characters might be considered as advanced or 
retarded, and it would be dangerous to take their occurrence as evidence of hybrid 
origin ; but the possibility of such effects needs to be considered in assessing the 
systematic importance of characters, and in tracing environmental correlations. 

Though these characters tend to appear together in forms of hybrid origin, the 
distribution of each is very erratic. The abrupt loss of black from the male tail 
(coinciding with a sharp increase in size, and the loss of ventral rufous) at the Ombai 
Strait, and its progressive reappearance eastwards here (H39 to 41) and in the 
Louisiades (H47a to c) suggests that this character may be dependent upon a rather 
precise balance between genes from each parental group. If this is true of all the 
" marks of hybridity ", and each has a different threshold value which depends also 
upon the parental genomes involved, the sporadic realization of these trends is not 
surprising. 

Sympatry 

All the forms of the pectoralis superspecies replace one another geographically, 
except in New Guinea. There P. soror and P. schlegelii represent one another 
altitudinally, but meet at about five thousand feet. There is no sign of intergradation 
between these very distinct species. They are the endemic representatives of P. 
pectoralis, and occupy the primary hill and mountain forest respectively. P. pectoralis 
itself has only been able to enter New Guinea by way of certain disturbed habitats. 
In the lowlands of the south and south-east E24~25a, it occupies coastal second 
growth, and may never come in contact with P. soror of the hill forest (though they 
are only three miles apart on Fergusson and Goodenough Islands respectively). 
Another race of P. pectoralis is known only from two river valleys on the northern 
slopes of the Snow Mountains E27. These valleys have been much affected by 
intensive native cultivation, which has stripped the forest from their floors and far 
up their sides. The local race of P. pectoralis seems to live largely in second-growth 
stands of Casuarina in the resultant grasslands, but also in the forest (Archbold, 
Rand & Brass, 1942). P. schlegelii descends a little way into the valley forest, and 
is found there side by side with P. pectoralis, with no signs of intergradation. Else- 
where in New Guinea P. soror is found up to the level of the valley floors, but it 
seems to be absent locally, and only appears in the hill forest more than a thousand 
feet lower. 

P. soror and the mountain race of P. pectoralis are so much more alike than 
intergrading forms outside New Guinea that one would expect that they must be 
conspecific. But the absence of interbreeding between them can scarcely be due solely 
to extrinsic barriers. They have been collected within a few miles of one another 
(on the only expedition which has yet encountered the montane race of P. pectoralis) 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 



167 



and must surely meet on occasion. P. pectoralis in the Snow Mountains is separated 
from its relatives in southern New Guinea by over three hundred miles of primary 
forest. It seems possible that further investigation will show the known populations 
to be a few of many, small and scattered among the mountains, wherever disturbance 
of the forest by man, landslide, wind, flood or fire provides a shifting foothold (cf. 
Rand, 1941, on the origin of the grassland avifauna). In any case, P. pectoralis 
has presumably passed through the range of P. soror to reach the known localities 
in the Snow Mountains. It is not strictly true to say (Rand, 1940) that this race of 
P. pectoralis shows no closer relationship to P. soror than does the southern New 
Guinea one. Discounting the effects in southern New Guinea E24 of gene-flow from 
group F, most of the changes (male throat feathers grey-based, collar reduced, mantle 
greener, edges of wing-quills olive and broader, female with olivaceous breast, brown 
cap and blacker tail) are in fact in the direction of P. soror. But even if these differences 





SCHLEGELII 

3 



Fig. 3. — The three sympatric forms, in the Snow Mountains of New Guinea. 
pectoralis E27 ; P. soror 2a ; P. schlegelii 3a. 



168 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

from other members of group E imply gene-flow from P. soror, rather than parallel 
adaptation to higher altitudes (p. 188), the introgression must have been slight and 
soon ended. The great reservoir of soror genes would otherwise soon swamp the small 
populations of P. pectoralis in the mountains, destroying not only the rather slight 
phenotypic differences but the genetic determination of habitat preference which 
presumably keeps the two forms apart. Although the evidence is very indirect, it 
seems that P. soror and group E of P. pectoralis are genetically sympatric species. 

A rather similar situation arises in the Bismarcks, where dahli E25 occupies 
small islands (but also occurs at certain coastal localities on the large islands) while 
the endemic race H48 keeps strictly to the dense forest inland (Dahl, 1899). Probably 
because of this difference in habitat preference, there seems to be little interbreeding 
between the two forms, though Hartert (1926) records that occasional males on New 
Britain combine the smaller size and greener mantles of the endemic race with the 
greyer wings of dahli, and Mayr (1955) suggests that much of the internal variation 
of dahli may be the result of gene-flow from group H. The two forms must meet 
occasionally, though dahli may not normally breed on the large islands. It would be 
reasonable to suppose that groups E and H must be partly isolated by intrinsic 
barriers, like P. soror and the Snow Mountains race of P. pectoralis. Yet in the 
Louisiades H47 the same groups have merged completely. 

Several explanations might be put forward to account for this anomaly, any or 
all of which may be true in part. It may be that in the Louisiades existing barriers 
have been broken down by hybridization (cf . Sibley, 1954ft) , permitting the populations 
to merge, while dahli has not been long enough in the Bismarcks for this to have 
happened. Or the populations of group H endemic on the relatively small islands of 
the Louisiades may have been inadequately shielded from interbreeding by differences 
in habitat-preference. Or after interbreeding in the Louisiades, dahli in the 
islands west of the Louisiades (E25D-C) may have secondarily developed barriers 
against group H, before invading the Bismarcks. In the absence of positive evidence, 
it is safest to assume that the two stocks in the Bismarcks represent one another 
geographically, and seldom meet in the breeding season. Yet it is conceivable that 
intrinsic barriers to interbreeding are effective here. From this, and from the different 
reactions of the schlegelii and soror assemblages (and of differentiated stocks within 
the latter) when they meet in New Guinea and in the archipelagos, it is clear that 
species-limits need not at all closely follow the pattern of descent and resemblance. 

P. schlegelii, P. soror and group E of P. pectoralis are genetically sympatric (Cain, 
1953) and must be regarded as good species. Yet the subspecies-groups A to H of 
P. pectoralis, some of them much less alike than are P. soror and group E, are inter- 
connected by a web of secondary intergradation (Text-fig. 4). In the absence of 
evidence that groups E and H meet in the breeding season without significant 
interbreeding, all these groups can be included in a single species. 

GROUP AFFINITIES 

The schlegelii assemblage 

P. schlegelii, groups A to D of P. pectoralis, and P. flavifrons all differ markedly 
from the standard patterns for the superspecies. Each is distinguished from the 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 



169 



others by characters which are not merely striking but stable and systematically 
important (as is shown by their co- variation) . At first sight they seem to be indepen- 
dently derived from more standard forms, rather than closely related to one another. 
There is little in common, for example, between P. schlegelii and the Fijian group D 
(Text-figs. 3 and 2, bottom). 

Yet there are important resemblances between members of this assemblage of 
forms. Most striking is the similarity between females of P. schlegelii and of P. 




Fig. 4, — Diagram of the phylogeny suggested for the superspecies, showing secondary 
intergradation between subspecies-groups of P. pectoralis (dotted) and sympatric 
co-existence in New Guinea (arrows). 



170 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 



pectoralis in the northern Moluccas (group B). Furthermore, the males of the same 
forms are alike in having black chins, and throat-patches overlapping the gorgets 
without black tips to the feathers. These two characters of the males are shared by 
forms in most of the groups of the schlegelii assemblage. The black chin is absent 
from all the hybrid forms and from group D, but characterizes P. schlegelii, P. flavi- 
frons and undiluted members of groups A to C. It is not found elsewhere in the 
superspecies (except in the black- throated race H57). The unusual structure of 
the gorget-edge is less affected by gene-flow, and is found in almost all the members 
of the assemblage which have gorgets. Elsewhere it occurs only on New Caledonia 
G34 (possibly related to the assemblage) and Ndeni H52 (see p. 162). 

Several other characters serve to link two or more groups. In this way each of 
the groups is united with at least one other : groups A to C of P. pectoralis to one 
another and to P. schlegelii, group D to group C, and P.flavifrons to group D. Table 
III sets out the main characters which occur in more than one group of the assemblage, 
while remaining rare elsewhere in the superspecies. The table does not illustrate the 
resemblance between females of P. schlegelii and of group B, since this depends on 
the conjunction of several characters (grey-barred throat, olive breast, grey auriculars 
and lack of phaeomelanins) which occur separately in several different forms outside 
the assemblage. 

Table III. — Characters Uniting the schlegelii Assemblage. 

schlegelii assemblage 





r 










" '"\ 


SU7UT 




schlegelii 


A 


B 


C 


D 


flavifrons , 


assemblage 


Male: 
















Chin black . 


. i-3 


1 


6-7 


0-15 


— 


X 


— 


Throat yellow 


— 


— 


— 


9-17 

(E26) 


18-21 


X 


H42 


No gorget . 


— 


— 


— 


12 


18-19 


X 


— 


gorget edge 


- i-3 


i-3 


6-8 

(H43) 


9-1 1, 
13-15 


— 


— 


. H52 


breast rufous 


• i-3 


i-4 


— 


17 
(G34) 


— 


— 


► — 


Forehead yellow . 


— 


— 


— 


— 


18-19, 


X 


— 


Mantle melanic 


— 


4 


— 


13-14 


2 1 

20 

(H55-56) 


X 


. H52 


Wing black 


i-3 


4 


— 


13-14 


— 


X 


— 


Tail black . 


i-3 


1 


6-7 


13 


— 


X 


H52 


Female : 
















Cheeks yellowish . 


— 


— 


— 


9, 11 & 12 


18-19 


— 


— 


Underparts streaked 


— 


— 


— 


9-16 


18-19 


— 


— 


Wing russet 


— 


— 


— 


9-16 


18-21 


— 


— 



The soror assemblage 

P. soror and the remaining groups of P. pectoralis are united negatively rather than 
positively, by the absence of those unusual characters which distinguish and link 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 171 

together the groups and species belonging to the schlegelii assemblage. Since group 
H as a whole scarcely departs from the average characters of the whole superspecies, 
it can be associated with other groups only negatively. However, there are a few 
characters and trends which link P. soror with groups E to G and these with one 
another, and are rather rare in the schlegelii assemblage. These are set out in Table 
IV. 

Table IV. — Characters Uniting the soror Assemblage (except Subspecies-group H). 









soror ■< 


assemblage 

A 




schlegelii 
assemblage 


Male : 


r 

soror 


E 




F 


G 


Carotenoid pale 
Quills grey 
Coverts yellow . 
Tail pale . 


• • 


i-3 

i-3 

i-3 


22 
22-25 
22-25 

22 




28-30 
28-30 
28-32 
28-32 


34. 36-38 

35 
34-37 


D19, 21 
. D18-19 
C17 


Female : 
















Throat white . 

Throat streaked (not belly) 

Cap grey (not auriculars) . 


i-3 


22-27 
25.27 
22-25 




28-33 


34-38 
38 


A4-5 
A 5 



Several of these characters seem to appear chiefly in higher latitudes, and may be 
related to the generally cooler climates encountered by these groups. Yet they are very 
different from the characters of P. schlegelii, which lives at higher altitudes than 
P. soror. The resemblances between group F and the Vanua Levuan race D19 
(grey-edged wings and yellow-edged wing-coverts in the male ; barred throat, 
melanic belly and almost complete absence of carotenoid in the female) may be due 
partly to selection under cooler climates. These characters, and others which (it 
may be objected) could be used to erect any number of alternative assemblages, 
are evidently unstable, since they appear independently in widely-scattered localities 
and cut across the discontinuities indicated by the co-variation of more stable 
characters. 

ARRANGEMENT 

The considerations of relationship, intergradation and sympatry discussed above 
permit the various forms to be arranged systematically. The categories of super- 
species, species, subspecies-group and subspecies are used, and considerable infra- 
subspecific geographical variation is recognized. 

The superspecies 

The category of Artenkreis or superspecies (Rensch, 1928; Mayr, 1931a) was 
introduced for groups of strictly allopatric representatives, some of which are too 
unlike to be considered conspecific. Mayr pointed out that the great practical 
usefulness of this category might be expected to decline as the possibility of unlike 
representatives being conspecific became generally accepted. More recently the scope 



172 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

of the superspecies has been extended to include groups of forms some of which are 
genetically sympatric (Cain, 1953), with their breeding ranges even overlapping 
slightly (Mayr and Vaurie, 1948 ; Mayr, 1949 ; and definition in Mayr, Linsley and 
Usinger, 1953). This extension gives the category permanent value, in expressing 
the relationship between forms which can never be considered conspecific yet which 
have barely ceased to represent one another geographically. But it removes the 
objective criterion which limited the use of the superspecies under the older usage, 
and opens the way to further extension. There is a danger of the category losing its 
special connotations of geographical replacement, becoming synonymous with the 
species-group. 

Sometimes a species or superspecies is represented locally by a pair of sympatric 
species, so that it would be arbitrary to include one rather than the other in the 
superspecies, or to exclude both. Cain (19546) has introduced the term " doublet " 
for such pairs in the Ptilinopus purpuratus superspecies, and this usage is helpful 
in indicating cases of double invasion. Where triple invasion by a single species has 
resulted in three representatives locally, the term " triplet " may be used. For 
example, Zosterops lateralis is represented by a doublet on Lord Howe (Z. strenua 
and Z. I. tephropleura), and by a triplet on Norfolk Island (Z. albogularis, Z. tenuirostris 
and Z. /. norfolciensis) . 

P. soror is so like many members of P. pectoralis, while P. schlegelii is so strikingly 
different, that the former is the obvious New Guinea representative. P. soror and 
P. pectoralis seem to be only genetically sympatric, without actual overlap, and might 
be included in one superspecies. But we have seen that, while P. soror is indeed the 
more closely related to some subspecies-groups of P. pectoralis, P. schlegelii is so to 
others. The two species should therefore be considered as a New Guinea doublet, 
like Ptilinopus coronulatus and Pt. pulchellus (Cain, 1954&). The local intrusion of 
P. pectoralis itself means that there is a triplet on the northern slopes of the Snow 
Mountains. 

Species 

All the remaining forms are geographical representatives. Those that intergrade 
must be conspecific, so that even the extreme forms on Sumba and in the northern 
Moluccas, Solomons and northern Fiji belong to P. pectoralis. Certain striking single 
characters have evidently arisen several times independently, and most modern 
systematists would agree in placing within P. pectoralis both the hen-feathered races 
on Rennell and Norfolk Island C16 & F33, and the melanic ones in the Solomons 
and Santa Cruz C13 '& H52. 

However, there are a few extreme forms (Text-fig. 5) whose status is doubtful, 
and can only be decided by comparing the degree of difference between them and 
their nearest relatives with that shown between sympatric species on the one hand, 
and between intergrading subspecies on the other. But in the pectoralis superspecies, 
two sympatric forms are much more alike than many intergrading ones : the 
relationship between visible differentiation and the establishment of barriers to 
interbreeding is not the same in New Guinea as in the archipelagos. The forms of 
doubtful status are all found on more or less isolated islands, whose avifaunas are 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 



173 



much poorer in species than those of New Guinea and include at most one other 
species of Pachycephala (P. rufiventris on New Caledonia) . It is reasonable to suppose 
that the intergradation of unlike subspecies in the archipelagos is more relevant here 
than the coexistence of like species in New Guinea (p. 179). 

P.flavifrons in Samoa is a derivative of the Fijian group D (Table III), but deserves 
the specific rank always accorded it. Group D is itself the most aberrant in P, 
pectoralis, and is probably on the borderline of ethological and genetic incompatibility 
with the rest of the species, while P.flavifrons has acquired further unusual characters 
— including the unique cock-feathering of its females. 

Previous authors have considered the Tongan form H57 as a distinct species 
(P. melanops) because of the black throat of the male. In other ways this form is 




FLAVIFRONS 





Fig. 5. Three unusual island forms. Samoa, P. 
H57 ; New Caledonia P. [p.] caledonica G34. 



lavifrons ; Tonga, P. p. melanops 



174 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

rather close to the standard pattern. It seems to be a member of group H, not more 
modified than might be expected from its isolation and the poverty of the avifauna 
in which it is placed. There is no reason to believe that the black throat reflects any 
very profound genetic change. The white throat-patch is very restricted on Sumba 
Ai and in P. schlegelii, and reduced to narrow white barring on black in individuals 
of the latter's close relative P. aurea. The mountain species P. nudigula and P. 
implicata (derived from the pectoralis superspecies) have acquired blackish throats 
independently. On the other hand, the black throat does alter the appearance of 
the male most markedly, and might therefore be of ethological importance. But so 
does the abolition of the gorget, and even more the total loss of the distinctive 
male pattern : yet there has been gene-flow between races with and without gorgets, 
while hen-feathered males in dimorphic races are able to find mates and breed. It 
is most improbable that the black throat alone would be sufficient to restrict inter- 
breeding while further barriers were selected for, in the event of a second invasion of 
Tonga. " P. melanops " ought therefore to be considered as a race of P. pectoralis. 

In the New Caledonian representative G34 also the male is unusual, the female less 
so. The male shows several unusual characters (long throat-feathers, rufous breast- 
patch and pale cap), which appear independently in other races. This form has been 
excluded from P. pectoralis, even by Mayr (1945), for nomenclatural reasons. Musci- 
capa caledonica Gmelin, 1789 — the valid name for it — antedates M. pectoralis 
Latham, 1801, so that merging them should mean changing the well-known name 
Pachycephala pectoralis to P. caledonica throughout its seventy-odd subspecies. It 
would be most undesirable to upset an established name in this way on the basis of 
a subjective decision. Yet it is also unsatisfactory to recognize the New Caledonian 
form, considerably less aberrant than the Fijian races, as a distinct species. Mayr 
(1941& and 1945) does not include P. caledonica in lists of the endemic species of New 
Caledonia. Since stability could be maintained by refusing to recognize the sub- 
specific status of caledonica, this is not a case which can be submitted to the Inter- 
national Commission for Zoological Nomenclature. I have avoided the dilemma by 
including caledonica among the subspecies of P. pectoralis, while enclosing the 
specific name in square brackets {Pachycephala [pectoralis] caledonica), an expedient 
suggested by Dr. A. J. Cain. 

The third form here included in P. pectoralis for the first time is the hen-feathered 
one on Salayer A5 (teysmanni). This has previously been considered as the repre- 
sentative, not of P. pectoralis, but of the hen-feathered species P. orpheus, which is 
very like females of group A. It agrees with the former species in its long bill, black 
in both sexes, and with the latter in its juvenile plumage (streaked below and without 
rufous on the wings), while it is intermediate in degree of dimorphism. Juveniles of 
the Sundan group A have less rufous wings than most in P. pectoralis, and a nestling 
from Pantar A3b is greyish with ventral streaking. P. orpheus occupies Timor and 
islands to the east, and it seems on the whole more probable that this and the Salayer 
form are independently derived from group A of P. pectoralis, than that P. orpheus 
has colonized Salayer across the three-hundred-mile gap occupied by P. pectoralis, 
and there partly reversed its loss of dimorphism. If this interpretation is correct, 
there is no reason to separate the Salayer Pachycephala from P. pectoralis. 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 175 

Subspecies-groups 

There is no formal category between the species and the subspecies, and I call the 
eight divisions of P. pectoralis subspecies-groups. Zeuner (1943) has used the term 
in a rather different sense. The recognition of subspecies-groups permits the inclusion 
of well-marked geographical representatives (e.g. Corvus cor one and C. comix : cf. 
Huxley, 1942, 249) in a single species, while still expressing both the distinction 
between them and the minor subspeciation within each. There are no nomenclatural 
rules for such informal categories, and it is convenient here to use . geographical 
designations rather than subspecies names for them. 

Subspecies 

No two populations have precisely the same distributions of genes, so that it is 
futile to attempt the subspecific separation of all local forms which differ statistically 
from the others. In avian systematics, some variant of the " seventy-five per cent 
rule " (see Mayr, Linsley and Usinger, 1953) is widely accepted as a lower limit for 
the constancy of differences between subspecies. This is a useful check on excessive 
splitting which should probably be retained whatever further considerations are 
introduced. But it does not take account of the magnitude and systematic importance 
of the differences. A very slight though constant difference in a single character 
which varies in response to climatic differences suffices to separate two forms ; while 
greater though rather less constant differences in several characters whose co- variation 
shows them to be largely independent of climate, and of systematic importance, do 
not. The subspecies-concept remains purely morphological, while that of the species 
now depends ultimately upon genetic isolation. 

In parts of the range of P. pectoralis, the birds from almost every island can be 
distinguished by careful comparison of long series. This is true especially of the New 
Hebrides (G36~37f) and the Bismarcks (E25d-j and H48a~49). Mayr (1945) recog- 
nized six subspecies in the New Hebrides, but now considers that most of these 
should be combined {in litt.) . The differences (almost entirely confined to the females) 
are slight, concern highly labile characters (such as the intensity of carotenoid and 
degree of phaeomelanization),intergrade through series of islands, and are exceeded 
by individual differences. The recognition of six subspecies conceals the true situation 
— that P. pectoralis is remarkably uniform throughout a great number of widely 
separated islands in the New Hebrides, though the population on Aneitum G36 is 
clearly distinguishable from the rest. I have therefore combined all the remaining 
populations (G37) . The Erromango population G37a deserves subspecific separation 
according to current usage (p. 205), but those from Efate to the Banks Islands 
G37b-f should be combined. 

Populations of groups E and H are distinguishable in series from island to island 
in the Bismarcks. The differences in group E are mainly in size, with populations of 
larger (E25b, c & d) and smaller (E25e) birds distributed apparently sporadically 
(cf. Cacatua galerita triton, Mayr, 1937). There are also slight differences in the 
pigmentation of the females and in the depth of carotenoid in the males. The 
situation is best expressed by combining the populations from south-eastern New 



176 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

Guinea to Nissan E25a-k in dahli — though that on Fergusson E25C is on the border- 
line of subspecific recognition by current usage. There is little greater variation in 
plumage between populations of group H in the Bismarcks H48a~49, and that little 
is almost entirely confined to the females. There has as yet been no attempt to 
divide the populations from Umboi to Lavongai H48a-c, although the females can 
be distinguished in series (Mayr, in litt.). Females on Mussau and Lihir H48d~49 
are more distinct — about as much so in relation to the other Bismarcks females as 
those on Erromango are in the New Hebrides. The Lihir form H49 is here separated 
subspecifically on account of its much greater size. But while that on Mussau H48d 
deserves separation under current usage, it is here combined with H48a-c to emphasize 
the homogeneity of this series of populations in contrast to those of Tabar and Manus 
H50-51, whose females are much more distinct. 

By considering these areas of incipient subspeciation, and raising the level of 
difference required for separation so as to clarify the pattern of variation, we have 
arrived at a pragmatic standard for the subspecies of P. pectoralis. When this is 
applied elsewhere in the species range, a number of named subspecies are combined 
with others. This is especially true of the "small-island forms " (p. 188), of which 
only those on Ternate and Tidore B7 and Lihir H49 are sufficiently distinct to be 
separated on size alone, at this level of difference. Some of the remainder (e.g. H43b) 
are distinct enough for separation according to the seventy-five per cent rule, while 
others (e.g. A3b) are perhaps not. 

The principle that minor subspecies should be combined, in order to emphasize 
the more important discontinuities, can be applied to variation on continents also. 
In Australia I have recognized only those forms considered by Mayr (1954a) to be 
original isolates — the subsequent expansion, intergradation and minor evolution of 
which have partly obscured the situation. Although the mangrove-living forms E22 
in north-western Australia are known only from a few widely-separated localities, 
their variation is wholly of clinal type. The situation could be expressed in nomen- 
clature as " P. p. cl. bynoei-melanura-violetae " (E22a-c-23), but it is convenient to 
separate the populations which show these regular changes from the more constant 
ones to the east E23. In P. schlegelii the populations of south-eastern New Guinea 
3b should be combined with those of the Snow Mountains 3a, because the changes 
are slight in comparison with those which distinguish the Vogelkop and Cyclops 
Mountain races A1-2, and intergrade through central New Guinea (Mayr & Gilliard, 
1954). In P. soror, on the other hand, there is a sharp change in the colour of the 
male tail in passing from the Bismarck and Saruwaged Mountains 2b to those of 
south-eastern New Guinea 3 — relatively important in this less variable species. 

The populations between Koro D2ob on the one hand, and Viti Levu and Vanua 
Levu Di8a & 19a on the other, are variable, and their hybrid indices evidently 
depend mainly on their distances from the parental populations. That on Taviuni 
D2oa is only slightly affected by gene-flow from Vanua Levu, and is best included 
with the Koro race. Those on Ovalau Di8b and Rambi and Kio D19C are also 
perhaps closer to the latter, but are considerably modified by gene-flow from Viti 
Levu and Vanua Levu respectively, and are therefore included in the subspecies of 
those islands. It does not seem desirable to recognize several hybrid races, differing 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 177 

only in hybrid index and variability (cf. Mayr, 1932b). If this were done, consistency 
would demand the recognition of the constant Santa Anna population 017b as 
subspecifically distinct from the variable ones on San Cristobal Ci7a. 

The subspecies numbered on Text-fig. 6, in the check-list, and in the text are 
delimited to make as clear as possible the pattern of isolation in the superspecies. 
It is not suggested that this arrangement should be generally adopted, since it does 
not accord with current usage, and a list of the subspecies to be recognized is therefore 
given on p. 205. But it does reveal the more important discontinuities which exces- 
sive splitting obscures, and the subspecies thus delimited have perhaps more 
biological significance, since some attempt has been made to consider discontinuity 
rather than mere constancy of difference (cf. Sibley, 1954a). It seems that the 
subspecies-concept might with advantage be modified by introducing such con- 
siderations. This would seldom require such considerable changes in the number and 
scope of subspecies as are necessary in the exceptionally variable species P. pectoralis. 



EVOLUTION 

Geographical speciation in New Guinea 

The distribution of species and subspecies in many New Guinea birds strongly 
suggests that northern New Guinea was once isolated (Mayr & Gilliard, 1952), and 
this agrees with what is known of the tectonic history of the island (Carey, 1938). 
Some of the northern endemics have since spread into the Vogelkop. The pattern 
of variation of P. schlegelii and P. soror and the distribution of forms related to them 
(discussed below) suggest that they were originally geographical representatives, 
with the former in the Vogelkop and northern New Guinea (P. schlegelii 1-2) and 
the latter in the rest of the island (P. soror 2-3). 

P. schlegelii has three distinct subspecies, with that of the Cyclops Mountains 2 
combining characters shown by those of the Vogelkop 1 and the Snow Mountains 3 
(and so perhaps ancestral to both) . The Vogelkop form has invaded the Wandammen 
Mountains, which suggests that the species was absent from the nearer Snow Moun- 
tains at the time of colonization. There is no sharp break between the Snow Mountains 
3a and the south-east 3b (though there are gradual clines), suggesting a rather 
recent expansion in this area. In P. soror, despite its greater overall homogeneity, 
there is a slight but definite break at this point (subspecies 2/3), while there is no 
appreciable difference between the birds of the Snow Mountains and of the north. 
This species has evidently colonized the northern mountains much more recently 
than those of the south-east. Both species have distinctly brown females in the 
Vogelkop, but in P. soror this is the only marked character of the Vogelkop race. 
Several birds are blacker in the Vogelkop than elsewhere in New Guinea, and this 
may be related to greater humidity (Cain, personal communication). If the parallel 
tendencies towards brownness in Pachycephala females here are adaptive, the changes 
may have been produced rather quickly. Also, isolation allows rapid change, 
unimpeded by gene-flow. P. soror may have been longer in the south-east than 
in the Vogelkop, despite the greater distinctiveness of the female in the latter area. 

Two species in New Guinea seem to be hen-feathered derivatives of P. schlegelii 



178 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

and P. soror. P. lorentzi is found within the geographical and altitudinal range of 
P. schlegelii in the Snow Mountains 3, and resembles the female of that species. P. 
meyeri bears the same relationship to P. soror in the Vogelkop 1. If these are relicts 
of early reciprocal invasions, their distribution tends to confirm that at one time 
P. schlegelii was absent from the Snow Mountains, and P. soror from the Vogelkop. 
Distribution and variation within P. pectoralis further reinforce this suggestion. 
Females of the Moluccan subspecies-group B, geographically near the Vogelkop, 
are remarkably like those of P. schlegelii (p. 169). Central members (subspecies- 
groups A to C) of the schlegelii assemblage are distributed towards the north of the 
species range, while the soror assemblage (especially the central groups E to G) is 
southerly. The early distribution of the assemblages according to this hypothesis is 
shown in Text-fig. 7, from which group H is omitted because its origins are obscure 

(p. 182). 



Origin of the dichotomy 

The presence of a doublet, representing locally a single more widespread species, 
implies double invasion. It has been suggested above that in the pectoralis super- 
species the second invasion took place within what is now New Guinea. If this is 
not true, the evidence suggests that P. schlegelii represents the endemic New Guinea 
stock, and that the soror stock arose as its geographical representative in Australia. 
While (outside New Guinea) the schlegelii assemblage is confined to islands, the 
soror assemblage occupies Australia. It must be rare for a form evolved in isolation 
on an archipelago, in competition with relatively few species, to compete successfully 
in the rich avifauna of primary forest on a continent, especially under the different 
physical conditions at high altitudes. But an Australian forest bird might well 
colonize the New Guinea hill forest. Gradual invasion across the Arafura shelf 
(often dry in the past) might favour the perfection of predeveloped isolating mecha- 
nisms (p. 180). The scattered and restricted subspecies-groups belonging to the 
schlegelii assemblage show a relict distribution comparable to that of the old species 
in the Rhipidura rufifrons superspecies (Mayr & Moynihan, 1946), while the soror 
assemblage occupies a wide and almost continuous range, like that of R. rufifrons 
itself, most of which has evidently been colonized comparatively recently. Patterns 
of intergradation in P. pectoralis, especially in the Banda Sea, suggest that the soror 
assemblage is still expanding (presumably as the result of selection) at the expense 
of the schlegelii assemblage. It is hard to accept the latter as a relatively recent 
colonist of New Guinea, in the face of competition from an entrenched P. soror. 

Since the intergradations of group C with the soror assemblage are evidently 
secondary (p. 160), it is necessary to pass through group H in order to derive P. soror 
from pvoto-schlegelii by way of primary intergradation in the archipelagos (see 
Text-fig. 7, right-hand inset of Text-fig. 6, and Text-fig. 8). Of the possible sequences 
by which it could be derived in this way, the least improbable is schlegelii-B-R-G- 
E+F — soror ; but it is not convincing. The almost inescapable conclusion is that 
the schlegelii stock arose from the common ancestor in New Guinea, and the soror 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 179 

stock either there or in Australia. This implies that P. schlegelii and P. soror are not 
merely terminal forms without issue, such as are produced by the double invasion 
of islands (e.g. Zosterops, p. 172 ; Ptilinopus mercieri and Pt. dupetithouarsii, Cain, 
19546), but represent the basal forms of their respective assemblages, geographically 
very near to their areas of origin. 

Since the sympatric species P. schlegelii and P. soror are connected by way of 
secondary intergradation in P. pectoralis, which proves to be specifically distinct 
from them both, this conclusion seems to imply reticulate evolution (Text-fig. 4). 
The origin of a genetically isolated new species from two existing ones is a commonplace 
in botanical systematics, but the mechanism commonly involved (allotetraploidy) 
seems to be incompatible with the genetic mechanisms of sexually-reproducing 
animals (see Dobzhansky, 1937). There is no question of any such mechanism being 
involved in the pectoralis superspecies, since the forms of P. pectoralis which are in 
the biospecific relationship to P. soror (E24 & 27) are pure members of the soror 
assemblage. However, the evolution of the superspecies appears to have been 
reticulate at the species level only if the present status of P. schlegelii and P. soror 
is conventionally reflected back in time to their point of divergence. At the time 
when gene-flow between proto-schlegelii and proto-sor or effectively ceased, permitting 
them to diverge in isolation, the barriers between them were probably almost 
entirely extrinsic ; and there is no reason to suppose that intrinsic isolating mecha- 
nisms had been perfected before they independently colonized the archipelagos and 
Australia. The existing mechanisms are local products of selection, and the supposi- 
tion that the primary dichotomy of the superspecies took place in New Guinea 
does not imply that established species barriers were later broken down. 



Development of isolating mechanisms 

While in New Guinea the schlegelii and soror assemblages are represented by 
distinct biospecies, in the archipelagos they have proved capable of interbreeding 
freely wherever they have met. Biologically, New Guinea is strikingly distinguished 
from the surrounding islands by its vastly richer fauna. It is the centre of distribution 
of Pachycephala, with twelve species (cf. Mayr, 1941a — P. aurea, P. lorentzi, P. 
meyeri, P. simplex (with griseiceps), P. hyperythra, P. modesta, P. rufwentris (with 
monacha), P. rufinucha, P. tenebrosa, and the pectoralis superspecies). At the time 
when schlegelii and soror were developing intrinsic barriers against interbreeding 
with one another, they were probably in effective contact with most, of the other 
nine. It may well be that the barriers were to some extent predeveloped before the 
two stocks met again, as a by-product of selection acting towards isolation from 
their sympatric congeners (especially P. meyeri and P. lorentzi). Similarly, P. 
soror' s barriers against the closely related subspecies-group E of P. pectoralis may 
have been prospectively developed, when the former came in contact with P. 
schlegelii. 

In Australia, P. pectoralis is in effective contact with P. simplex, P. nifiventris, 
P. lanioides and P. olivacea (of which the first two are probably rather recent arrivals 

ZOOL. 4, 4. 13 



180 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

— Mayr, 1954a). Possibly the specific distinctness of group E from P. soror is partly 
related to the development of barriers against these, and especially against the 
northern mangrove species P. lanioides and P. simplex. 

In the archipelagos P. pectoralis meets with no more than one congener on any 
island : with P. grisola on Java and Bali (A2) ; with P. nudigula on Sumbawa 
and Flores (A3a) ; with P. orpheus on Timor and Wetar (H39) ; with P. rufiventris 
(including the griseonota group) on Damar, Tenimber, the Moluccas and Sula Isles, 
Rossel and New Caledonia (H41-45, B6-8, H47C & G34) ; with P. simplex on Fergus- 
son (E25C) ; and with P. implicata on Bougainville and Guadalcanal (C9 & n). Of 
these P. grisola in the lowlands, and P. nudigula and P. implicata in the mountains, 
are altitudinally separated from P. pectoralis, while P. rufiventris probably occupies 
drier habitats on the average (as in Australia) and shows altitudinal separation on 
Ceram and Buru (pp. 189-190). Only on Timor and Wetar is there an overlap, of long 
standing and without marked ecological separation, between P. pectoralis and a 
closely-related species, P. orpheus. On Timor and Wetar, P. pectoralis is large in 
comparison with its relatives to the west, and with sympatric P. orpheus ; which in 
turn has large representatives (par) on Roma, Letti and Moa, where the former does 
not occur. This suggests that there has been selection for size difference between 
the two species in the zone of overlap, as between Sitta neumayer and 5. tephronota 
(Vaurie, 1950). It is noteworthy that in this region the intergradation between 
subspecies-groups of P. pectoralis is strongly stepped at the Ombai Strait (between 
subspecies-groups A and H), which suggests that gene-flow has not been as free as 
in the Moluccas and Fiji. 

It seems therefore that the predevelopment of isolating mechanisms between the 
schlegelii and soror assemblages may be related to the intensity of selection against 
interbreeding with related species, to which the populations concerned were subject 
before they met. The degree to which they had developed different ecological 
preferences may also be relevant. It is probably more than a coincidence that group 
E of P. pectoralis, which is genetically isolated from its close relative P. soror in 
New Guinea (and to some extent from group H in the Bismarcks), is the most 
ecologically specialized in the superspecies (p. 190). New Guinea provides greater 
opportunities for altitudinal separation than any of the surrounding islands, and 
P. schlegelii and P. soror replace one another in this way to an extent seen in the 
archipelagos between P. pectoralis and much less closely related species. 

Another factor which may have affected the reaction of the two stocks on meeting 
is the manner in which this took place. Within New Guinea they would at first have 
been opposed to one another over a broad front, and separated by the relatively 
unsuitable habitat of lowland forest. Invasion through this would be slight but 
continuous, producing rare hybrids in the lowland zone. These would be at a selective 
disadvantage, and at the edge of this zone there would be strong selection in favour 
of isolating mechanisms. Once developed these would allow the stocks to overlap, 
and the developed mechanisms would spread back into the populations, followed by 
waves of reciprocal invasion. The same would happen where group E of P. pectoralis 
met P. soror at the edge of its range, and the development of isolating mechanisms 
would finally permit the former to penetrate the range of the latter as a distinct 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 181 

biospecies. In the archipelagos, on the other hand, the second colonizing wave 
(represented largely by group H) would have arrived over water, and its genetic 
representation in the population would probably be increased less by repeated 
invasion than by selection. In the absence of effective pre-established isolating 
mechanisms the new element would be incorporated in the endemic population, 
and no barriers could subsequently be developed. 

A suggested course of events 

Several alternative sequences of invasion and differentiation might be suggested, 
any of which could have produced the existing pattern of variation. It would not 
be profitable to discuss at length the pros and cons of these hypothetical alternatives, 
since the evidence does not seem adequate to decide between them with any finality. 
However, the following sequence seems to fit the facts more neatly than any alterna- 
tive, and is perhaps worth putting forward for comparison with those deduced for 
other groups in the area (e.g. Ptilinopus purpuratus species-group in Ripley & 
Birkhead, 1942; Halcyon chloris species-group in Mayr, 1949; Coracina species-groups 
in Ripley, 1941, and Voous & van Marie, 1949; Rhipidura rufifrons species-group in 
Mayr & Moynihan, 1946 ; Dicrurus hottentottus superspecies in Mayr & Vaurie, 1948; 
Dicaeum cruentatum-hirundinaceum species-group in Mayr & Amadon, 1947). It is 
presented without qualification, for the sake of brevity. 

It seems that the common stock of the superspecies arose in New Guinea (where 
the greatest concentration of Pachycephala species is to be found at present). P. 
nudigula on Sumbawa and Flores and P. implicata on Bougainville and Guadalcanal 
probably represent the relicts of an early burst of colonization westwards and 
eastwards respectively. The north-western and south-eastern populations in New 
Guinea, more or less effectively isolated from one another, diverged as subspecies. 
From the west and north " schlegelii " colonized the Lesser Sunda Isles (proving 
specifically distinct from P. nudigula), Moluccas, and Solomons (proving specifically 
distinct from P. implicata). The resulting widely-separated populations became 
very different from one another. Internal diversity may have been developed in 
groups A and B to much the same extent as is seen in group C to-day, before it was 
obscured by gene-flow. New Caledonia was probably colonized from the Solomons 
before the development of the special characters of group C, while there was a later 
expansion to Fiji and on to Samoa. Meanwhile the " soy or " subspecies had colonized 
Australia. Because of the more humid climate and frequent emergence of the Arafura 
shelf during the Pleistocene, its range was probably more or less continuous. Not 
until the colonization of Southern Melanesia and the separation of northern and 
southern populations by the Recent eremiation of Australia (Browne, 1945) could 
the internal differentiation of the soror assemblage proceed far. 

When the two New Guinea forms invaded one another's ranges as P. schlegelii 
and P. soror (their hen-feathered representatives P. lorentzi and P. meyeri having 
already done so), a contemporary systematist would have expressed the situation 
in terms of a superspecies (the schlegelii assemblage) overlapping in New Guinea 
with a polytypic species (the soror assemblage) . Yet the assemblages fused when they 



182 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

met in the archipelagos, mainly as a result of the explosive expansion of group H 
(black in right-hand inset, Text-fig. 6). Probably the complex palimpsest produced 
by new colonists interbreeding with the old, and incorporating characters proper to 
them, appeared first in New Caledonia at the invasion of Southern Melanesia from 
Australia. 

The origin of group H is obscure. The most distinctive forms belonging to this 
group are widely separated (p. 160), and so do not help in determining the direction of 
expansion. Its derivation from one of the groups belonging to the schlegelii assemblage 
would imply too much reversal and convergence to be plausible. Since its common 
characters are very close to the standard for the whole superspecies, it may even 
represent a rather conservative though highly adaptable derivative of the common 
ancestor in New Guinea. Or the resemblance between its Banda Sea and Pacific 
sections may be convergent. But on the whole group H seems most likely to have 
arisen from the soror assemblage, as a stock adapted to island life, in the Banda Sea. 
Former island populations between Tenimber H42 and the Louisiades H47 may have 
become extinct during the Pleistocene fluctuations in sea-level over the Arafura 
shelf. Whatever its origin, the group has spread widely until its influence is apparent 
from Java to Tonga, and has interbred with very unlike island forms belonging to 
both assemblages. At this time the contemporary systematist would have regarded 
P. soror as conspecific with the whole complex of intergrading forms, and would 
have looked on the superspecies as analogous to a ring species — with P. soror and 
P. schlegelii specifically distinct in New Guinea, yet connected through series of 
interbreeding forms in the archipelagos. The latest major event, the escape of group 
E from Australia as a colonist of coastal and second-growth formations, shows that 
this is an oversimplification, and that morphological analogy may be misleading in 
the assessment of potential isolating mechanisms. Group E penetrated the range of 
P. soror as a distinct, though very similar and closely-related, species ; merged 
completely with its near relatives of group H in the Louisiades, yet remains more or 
less isolated from the same group in the Bismarcks ; and has interbred freely with 
the very dissimilar and phylogenetically distant group C in the Solomons. 



Rate of divergence 

In general, the degree of difference now to be seen between related forms (both 
within and between subspecies-groups) agrees reasonably well with the sequence of 
events suggested above, when the retarding effect of gene-flow between populations 
has been taken into account. The groups belonging to the schlegelii assemblage 
are much more different from one another than those of the soror assemblage. Group 
C shows the most internal differentiation of any group in the superspecies, while there 
is considerable variation within group A despite the levelling effect of gene-flow 
from group H. Group D is evidently derived from group C, and there is remarkably 
little difference between the populations on Viti Levu and Vanua Levu ; while 
P. flavifrons, derived from group C at second hand, shows no appreciable variation 
between Savaii and Upolu. The females of group B are so like those of P. schlegelii 
that the populations in the northern Moluccas may have been in genetic contact 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 183 

with those of the Vogelkop long after the invasion of the Lesser Sunda Isles and 
Solomons, and the slightness of the geographical variation there bears out this 
supposition. 

P. schlegelii and P. soy or are here supposed to be New Guinea endemics of equal 
antiquity, and the former to have occupied a discontinuous range before their 
reciprocal invasions — so that its greater geographical variation is not unexpected. 
Similarly, the comparative uniformity of the Australian groups agrees with what is 
known about the rates of divergence in populations of continental and of insular 
range. In the characters common to most of its members, group G is little more 
distinct from groups E and F than they are from one another, and combines characters 
of each (Table IV) . This suggests that southern Melanesia may have been colonized 
at about the time that increasing aridity effectively divided the range of P. pectoralis 
in Australia. However, there is considerable variation within group G (even when the 
effects of a genetic contribution from the schlegelii assemblage to New Caledonia 
have been allowed for) and the much greater uniformity of groups E and F must 
be attributed to their more or less continuous continental ranges. Both these groups 
have isolated representative populations on small islands far from Australia, but 
hen-feathering on Norfolk Island F33 is the only considerable change shown by these 
populations. The distribution of group E in New Guinea and the nearby islands 
strongly suggests the very recent incursion of an ecologically-specialized form, and 
recent colonization is the probable explanation for lack of divergence on Lord Howe 
F32 also. 

Although there is considerable local variation within group H, widely separated 
forms are remarkably similar (for example, the females in the southern Moluccas, 
Bismarcks and Santa Cruz H43-44, 48-49 & 52-53). Apart from the effects of 
intergradation with other groups, the isolated sections of this group in the Banda 
Sea, eastern Papuan islands, Santa Cruz and Fijian archipelagos are not well 
differentiated from one another. It seems probable that most of the large range 
of this group has been colonized relatively recently, despite the wide scatter of very 
distinctive forms (p. 160). This conspicuous divergence of some of its members, in 
one or a few characters, is discussed below (p. 184). 

Unexpected uniformity 

In some areas there is surprisingly little geographical variation from island to 
island, although the distances involved are not small in comparison with those 
between islands occupied by strikingly different forms in other archipelagos. Where 
there is evidence of gene-flow between groups (as in the Banda Sea, Louisiades and 
Fiji), it is evident that the populations on different islands are or have been in genetic 
continuity, and the uniformity may be attributable to swamping. Elsewhere it may 
reasonably be explained in terms of colonization too recent for much subsequent 
diversification, as has been suggested for group H as a whole, and for the insular 
expansions of groups E and F. 

But the slightness of the geographical variation within the New Hebrides G36~37f , 
in contrast to the marked differences between the forms on the different island- 



184 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

groups of Southern Melanesia (G34, 35, 36 + 37, & 38), does not at first sight seem 
to be susceptible to either of these explanations. There is no evidence for exceptional 
gene-flow here (except perhaps from New Caledonia G34 to Aneitum G36), and the 
New Hebridean type is too distinct from its relatives for the whole archipelago to 
have been colonized only recently. The distances between islands in the New 
Hebrides are shorter than those separating them from the Loyalty Isles G35 and 
Vanikoro G38, but there are several gaps (e.g. from Erromango to Efate G37a/b, 
and from Maewo to Gaua G^ye/i) comparable with those which separate very distinct 
forms on Guadalcanal Cn, Malaita C12 and San Cristobal C17, or on New Caledonia 
G34 and the Loyalty Isles G35. Almost all the New Hebridean gaps are wider than 
those which separate the three marked races of the central Solomons C13-15. 

Much of the New Hebridean avifauna shows this combination of a fairly high 
degree of endemism with surprisingly slight geographical variation. This might 
follow if geographical variation were closely related to environmental differences, 
and if the New Hebrides were environmentally much more uniform than the other 
archipelagos. But, in Pachycephala at least, variation does not seem to be as minutely 
correlated with demonstrable local differences as this theory would demand, while 
the New Hebrides show much more variation in climate and vegetation (from a 
marked dry season, with permanent grasslands, in the south to a very equable 
equatorial climate, with only rain forest, in the north) than do the Solomons. It 
seems that for a considerable part of the avifauna, genetic isolation between the 
islands of the New Hebrides is either imperfect or only recently established. The 
existing patterns of variation might result from the splitting up of an originally 
continuous range (by subsidence), or from an expansion into new territory (by 
island-building or other external changes, since several species are involved). 

The geological evidence (Mawson, 1905) is that most of the New Hebridean 
islands except Malekula and Santo G37d have arisen since the late Pliocene, either 
by volcanic extrusion or by uplift. The terraced profile of parts of the central New 
Hebrides is very marked, and quite unlike anything seen in the eastern Solomons 
(Cain & Galbraith, personal observation). It is possible that geological events, and 
their ecological consequences, can be invoked in partial explanation of the avifaunal 
peculiarities of San Cristobal, including the incursion of a Southern Melanesian 
Pachycephala (Galbraith, in preparation). 

Unexpected diversity 

Some forms, on the other hand, are more different from their close relatives than 
is to be expected from the suggested sequence of events on the assumption that 
populations not swamped by gene-flow have diverged at approximately the same 
rate. The outstanding forms (i.e. A4 & 5 ; C12, 13, 14, 16 & 17 ; D21 ; E27 ; F33; H42, 
52 & 57 ; P.flavifrons (an offshoot of group D), and the subspecies (1) of P. schlegelii 
and P. soror) occupy more or less restricted ranges, isolated at the periphery of their 
respective groups. The law of peripherally-isolated populations (Mayr, 1954b) is 
well illustrated in areas where geographical variation is less advanced : the compara- 
tively well-marked forms in the northern Moluccas (By), New Hebrides (G36, & G37a) 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 185 

and Bismarck archipelago (H49-51, & H480!) are all peripheral. Mayr discusses the 
possible explanations : drift, resulting from small population size ; selection, 
resulting from environmental differences (physical and biotic) ; and " genetic 
revolution ", resulting from a small gene-pool (initially because of the smallness 
of the founding population, and subsequently because replenishment by gene-flow 
is restricted) . 

Because such aberrant peripherally-isolated populations often occupy relatively 
small islands, drift has been invoked to explain them. But few forms of P. pectoralis 
are found on tiny islets (the exception, group E, has as yet diverged little over a 
widely scattered range), and it is improbable that the local effective breeding popula- 
tions of this common bird are sufficiently small for selection to be overcome by 
random fixation and elimination of genes. 

The brownness of the females of both P. schlegelii and P. soror in the Vogelkop 
may be an adaptation to very humid conditions (p. 177), while there is evidence of 
the selective influence of climate on P. pectoralis, not only in Australia but to some 
extent in the archipelagos (p. 187). Certain regularities of geographical variation in 
different bird species suggest that the physical environment varies from island to 
island more than is generally supposed, and considerable local differences in climate 
between habitats within one island support this view (Cain, unpublished) . Unfortu- 
nately, meteorological data for this region are scanty, and are probably affected less 
by the relatively slight changes in average climate from island to island than by the 
precise siting of the stations. Although differences in the physical environment 
cannot be ruled out, the diversity of characters involved makes it necessary to look 
further for the causes of differentiation in peripherally-isolated forms. 

Biotic differences (especially in the avifauna) seem more promising, but are still 
more difficult to assess. Trends in the dimensions of forms on small islands are 
discussed below (p. 188), and are probably related to ecological redeployment. Most 
of the divergent forms occur in avifaunas which are relatively poor in species, and 
most of the variations in male plumage reduce the distinctiveness of the pattern. 
But the two features of the avifauna which are most likely to impinge on the visual 
properties of the pattern — the presence or absence of visual predators, and of species 
with similar patterns — do not provide a comprehensive explanation of the variation. 
Lack of visual predation might be expected to relax dorsal crypsis, and in fact two 
of the the three forms with black mantles are restricted to islands without known 
hawks (Samoa (P. flavifrons), and Ndeni and Santa Cruz islets H52). But both 
Accipiter novaehollandiae and A. albogularis are found on Vella Lavella together 
with the black-mantled Pachycephala C13, and the latter hawk may have been over- 
looked on Ndeni. Only in Australia and New Guinea, the Lesser Sunda Isles and New 
Caledonia does P. pectoralis overlap with other Pachycephala species having similar 
male patterns (P. soror, P. schlegelii, P. aurea, P. rufiventris and P. lanioides). The 
loss of the gorget on Malaita C 12, for example, is not solely related to the absence of 
such species, since the only other Pachycephala in the Solomons (P. implicata) is 
restricted to the mountains of Bougainville and Guadalcanal and does not have a 
conspicuous pattern. But such a loss might be disadvantageous in a richer avifauna 
containing species of similar plumage and behaviour patterns. 



186 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

Mayr (i 954ft) has pointed out that when an island is colonized by a few stray 
pairs from a large and genetically variable population, the initial representation of 
genes will be more or less random . The selective changes involved in regaining a 
coadapted genetic system will depend on the genes available, so that unpredictable 
differences between isolated populations arising in this way are to be expected. 
However, in all these populations there will be selection in favour of genes which 
have a favourable effect in the homozygote and are at an advantage against a more 
uniform genetic background, so that some regular trends may be looked for. Both 
these expectations are fulfilled in P. pectoralis. Most of the variations seem to appear 
entirely at random, yet the ones which strikingly affect the pattern occur only in 
such peripheral localities. Melanism of the mantle is confined to more or less isolated 
forms on rather small islands (Djampea, Kalao tua and Madu, Vella Lavella, Ganonga, 
Rendova and Tetipari, Ndeni and the Santa Cruz islets, Taviuni, Koro, Ngau, Ongea 
Levu, Fulanga and Wangava, and Upolu and Savaii). Furthermore, the traces of 
gene-exchange between populations separated by several miles of sea are unusually 
clear in P. pectoralis, and the rapid falling-off in gene-flow with increasing distance 
will obviously have important genetic effects in this species. 



ADAPTATION 

Character and climate 

The adaptive significance of geographical variation in relation to climatic differences 
emerges most clearly in continental areas, exposed to regular climatic gradients and 
occupied by continuous populations. Irregular differences in climate, and the effects 
of isolation and bursts of colonization by different stocks, make it difficult to correlate 
character and climate in insular regions (cf. Snow, 1954). In many species of birds, 
clinal changes are strongly marked from north to south down the eastern coast of 
Australia, in relation to decreasing average temperature. In this area P. pectoralis 
presents the appearance of a series of isolates with secondary intergradation, rather 
than of a continuous cline (Mayr, 1954a). The Tasmanian population F29 is isolated 
by sea, and there seems to be a gap in the range of the species between southern 
Queensland F3ia and the rain-forests around Cairns F3ib. There is no reason to 
doubt that the populations from eastern Victoria to southern Queensland F3oa~3ia 
are continuous, and material from intermediate localities may show the phenotypic 
changes to be gradual. 

We may arrange the three presumptive isolates of the east coast in a southwards 
series ; and since groups E and F are closely related we may perhaps add the Northern 
Territory isolate as a first member. If the northern Queensland populations F3ib 
were included, they would introduce a slight reversal of the otherwise progressive 
changes. There has evidently been gene-flow from this region into Cape York and 
across the Torres Straits, and it seems possible that reciprocal flow has produced a 
retardation of the female characters here (p. 166). Therefore the southern Queensland 
form F3ia instead will be taken to represent group F in the north of its range. Our 
series from north to south is then : (a) Northern Territory E23, (b) southern Queens- 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 187 

land F3ia, (c) eastern Victoria F30a, (d) Tasmania F29. In this series the following 
progressive (though neither smooth nor synchronous) changes are apparent : 
(i) increasing wing-length (and presumably body-size) ; 
(ii) increasing relative tail-length ; 
(iii) decreasing absolute and relative bill-size ; 
(iv) decreasing amount of solid black (in the male tail) ; 

(v) decreasing concentration of carotenoid (female plumage and male tail 
especially) . 

Such a series of progressive changes might be maintained without climatic adapta- 
tion, by gene-flow from end-forms which had diverged in isolation. The characters 
of an intermediate population would then depend mainly on its degree of genetic 
isolation from each gene-source. However, changes (i), (iii) and (iv) (increasing 
body size, decreasing relative length of appendages and concentration of melanins) 
are sufficiently general among homoiothermous animals, in relation to increasing 
latitude, to be recognized as Bergmann's, Allen's and Gloger's ecological rules (see 
Huxley, 1942). Furthermore, all five of these changes are paralleled in Palaearctic 
titmice (Parus spp., Snow, 1954), in relation to decreasing temperature. It is most 
probable that the changes seen in P. pectoralis in eastern Australia are likewise 
adaptive. In certain other parts of the species-range parallel trends are discernible, 
although (as in Parus) there are numerous exceptions among island forms. 

From the Northern Territory to midwestern Australia (E23~22a) , solid black and 
intensity of diffuse melanin and of carotenoid decrease, but wing-length also decreases 
and proportions are not much affected. The small size and general pallor of these 
forms are paralleled in group A of the Lesser Sunda Isles, and it is noteworthy that 
these two areas are the driest occupied by the superspecies. In many groups of 
birds (Snow, 1954 ; Cain, unpublished), intensity of pigmentation falls off with 
decrease both in temperature and in humidity. This applies both to melanins 
(Gloger's rule) and to carotenoids. In arid regions the range of temperature is great, 
and it may be that coloration and body-size are responding to lower minimum and 
higher maximum temperatures respectively. 

In group G, the male tail is black in the north G38, olive in the south G34-36, and 
olive with black subterminally in the centre of the range G37. There is a southwards 
decrease in the intensity of carotenoid, especially in the females, from Vanikoro 
G38 through the New Hebrides as a whole G36-37 (with local variation) to New 
Caledonia G34 ; but the Loyalty Isles race G35 is yellowest of all, and no correspond- 
ing regularities in the variation of size and proportions are apparent. Throughout 
the archipelagos there are such hints of climatic correlations, but the exceptions are 
so numerous that they cannot be relied upon. Thus at the level of the subspecies- 
groups within the schlegelii assemblage, it might be suggested that the low intensity 
of carotenoid in the females of groups A and D is correlated with the aridity of the 
Lesser Sunda Isles and the coolness of Fiji respectively ; while in the hot and moist 
Moluccas females of group B have intense yellow pigment. But there is as great 
variation in this character between adjacent islands of the Solomons (C9, 11 & 12 
against 10 & 13-17), whose climates cannot be very different. Another possible 
climatic correlation, since both localities are rather arid, is in the appearance of a pinkish 



1 88 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

phaeomelanic wash over breast and belly of the females in southern Australia F28 
and on Timor H39. But the appearance of the character also on small islands of the 
Flores Sea A4, and its absence from females of group E from the most arid parts of 
Australia, seem to contradict this. As in Palaearctic tits, climatic correlations are 
merely hinted at among isolated forms. 

Character and habitat 

In New Guinea members of the superspecies occupy different altitudinal belts, 
with more or less overlap, from coastal second-growth (P. pectoralis E24) through 
hill forest (P. soror) and lower montane cleared land (P. pectoralis E27) to mountain 
forest (P. schlegelii) . The three highland forms of this series agree in the olivaceous 
wash on the underparts of their females — found elsewhere in the superspecies only 
in forms derived from P. schlegelii. The relict species P. nudigitla and P. implicata 
(in the mountains of the Lesser Sunda Isles and Solomons) also show this character, 
which thus seems to have been independently acquired about five times by members 
and close relatives of the pectoralis superspecies, in relation to life at higher altitudes. 
The males also of P. nudigula and P. implicata are strongly washed with olive 
beneath, while those of P. soror have an olivaceous appearance (due largely to the long 
grey feather bases showing through). Possibly the rufous ventral wash of male 
P. schlegelii represents a parallel adaptation to high altitudes. The small size of the 
white throat-patch in P. schlegelii and P. soror, the reappearance of grey bases to 
the white feathers in P. pectoralis E27, and the greying or blackening of the throat in 
P. nudigula and P. implicata may also be parallel responses to similar environments. 
All these trends are towards the blurring of the conspicuous ventral pattern (obscured 
also in the montane species P. rufinucha, P. tenebrosa and P. olivacea). 

Several subspecies have been described from the Banda Sea, which inhabit small 
islands and closely resemble those of adjacent large islands, but which are described 
as being larger, with conspicuously larger bills and often with more golden-olive 
mantles. This is true of Lomblen, Pantar and Alor A3b compared with Sumbawa 
and Flores A3a ; Wetar H39b compared with Timor 1139a ; Amboina H43b 
compared with Ceram 1143a ; and Ternate and Tidore B7 compared with Morotai, 
Halmahera and Bat j an B6. Unfortunately the series of several of these forms 
available to the authors of the descriptions and to me are not adequate for the 
statistical significance of the differences to be determined. In the Bismarck archi- 
pelago the populations on Lihir and Tabar H49-50 are conspicuously larger and larger- 
billed than those of the large islands H48 & 51. The same is true of dahli E25 in 
comparison with its close relatives in northern Australia E23 (the populations E25 
lying between, being affected by gene-flow, are ineligible for comparison) and of the 
races on Lord Howe and Norfolk Islands F32-33 in comparison with the parental 
populations at the same latitudes in Australia F3ia. All the races which are strikingly 
larger than their close relatives (and have correspondingly large bills) occupy small 
or low-lying islands : Ternate and Tidore B7, the Loyalty Isles G35, the Tenimber 
Isles H42, Lihir and Tabar H49 & 50, and Tonga H57. This tendency towards large 
size, and particularly towards large bill size, is well known among insular birds 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 189 

(Murphy 1938 ; Mayr & Vaurie 1948), and may be related to redeployment of the 
few species on such islands among the available food niches. However, Mayr's 
figures (1932a, b) do not suggest any such general rule in the Solomons, Santa Cruz, 
New Hebrides and Fiji, for situations where closely related populations occupy 
neighbouring islands of different sizes. 

It has already been noted that markedly divergent subspecies occur on rather 
isolated, and usually small, islands. Apart from the effects of isolation and the small 
gene-pools of founding populations, the relative paucity of species on these islands may 
have been an important factor in permitting the male pattern to diverge widely 
from the standard. For example, in group A hen-feathering occurs on Salayer A5 
(about 35 resident land bird species) and melanism of the mantle on islands in the 
Flores Sea (about 55 species), whereas the male on Sumba Ai (about no species) 
is ventrally conspicuous and dorsally cryptic. In group C hen-feathering occurs on 
Rennell C16 (about 35 species) and melanism on peripheral islands of the central 
Solomons C13-14 (about 60 species), while the male on Guadalcanal Cn (about 95 
species) is standard for the group. The male on Norfolk Island F33 (about 15 species) 
is hen-feathered, unlike its relatives in eastern Australia F30-31 (several hundred 
species). The mantle is wholly black on Ndeni and the Santa Cruz islets H52 (about 
20 and 10 species respectively), and in P. flavifrons of Samoa (about 30 species). 
In Fiji the correlation with avifaunal poverty is less clear : black-mottled mantles 
occur on Koro, Ngau and the southern Lau archipelago D2ob, H55-56 (25-30 species) 
and not on Viti Levu (about 50 species) ; but on two islands with apparently similar 
avifaunas (about 40 species), black mottling appears on Taviuni D2oa but not on 
Vanua Levu Di9a. It may be that further species remain to be discovered on Vanua 
Levu, a large and mountainous island, whereas Taviuni is smaller and better known. 
But elsewhere, too, the realization of these trends seems capricious. Although neither 
hen-feathering nor melanism of the mantle occur where the avifauna is rich, they 
are not always present where it is poor. Well-marked races which do not show 
these tendencies occur on the following small, low or isolated islands : Tidore and 
Ternate By, Russel Islands Cioc, Vatu vara D21, Lord Howe F32, Loyalty Islands 
G35, Aneitum G36, Vanikoro G38, Wetar, Babar, Damar and Tenimber H39D-42, 
Lihir, Tabar and Manus H49-51, Utupua H53, Kandavu H54 and Tonga H57. 

Variation in habitat 

The members of the pectoralis superspecies throughout its range are forest birds 
which forage for soft-bodied insects (and some berries) among the twigs and branches 
of the substage and lower canopy. Field notes are rare in the literature, but it is 
clear that there is considerable variation in the habitats selected by different forms. 
The altitudinal deployment in New Guinea has already been mentioned, and races 
of P. pectoralis are found at different altitudes on different islands. 

Group A is found only above 6,500 ft. in eastern Java A2a and above 3,000 ft. 
on Bali A2b, but from 4,000 ft. down to sea-level on Flores A3a (Hartert, 1897 ; 
Stresemann, 1913 ; Meise, 1929 ; Rensch, 1931 ; Hoogerwerf, 1948). This may be 
related to the presence of the lowland P. grisola on Java and Bali, and of the moun- 
tain P. nudigula on Flores. 



igo THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

Altitudinal preferences are known to differ from island to island in the Solomons, 
though the details remain to be worked out. Mayr (1932a) reports the species as 
rare or absent in the lowlands of Bougainville and Malaita Co, & 12, though common 
near the coast on Choiseul Cioa. Cain and Galbraith (1956) heard it occasionally 
in the lowland forest on Guadalcanal Cn, but found it much more common in the 
hill forest at about 2,000 ft., and up to the lower limit of the mist forest (where it is 
replaced by P. implicata). On San Cristobal C17 I found it to be common at 2,000 ft. 
and down to the coast. It is recorded by Donaghho (1950) in the lowland forest on 
Guadalcanal, but not by Sibley (1951) in the same habitat on New Georgia C15. 

The altitudinal preferences of P. pectoralis in the northern Moluccas and Fiji do 
not seem to have been recorded. P.flavifrons is found at 600 ft. and above on Upolu 
(Nicoll, 1904), 

On Timor H39a, P. pectoralis occupies a greater vertical range than has been 
recorded elsewhere, from sea level to 7,500 ft. (Stein, 1936 ; Mayr, 1944c). It is 
absent from the lowlands (occupied by P. rufiventris) on Ceram 1143a and Buru 
H44, and reaches 5,000 ft. (Stresemann, 1914a, b). Scott (1946) records it on Santo 
G37d as commonest in the higher forest, though frequent also in the open lowland 
forest. I have seen it on that island and on Efate G37b in tangled second growth 
near the shore, a habitat never seen to be occupied by the species on Guadalcanal and 
San Cristobal. In southern and eastern Australia it is found in dense and open 
forest (records in the Emu, 1902 to date), but not in the dense myrtle-beech forest 
of the wettest areas on Tasmania F29 (Lawrence, 1952), where P. olivacea is found. 

The most striking specialization in habitat is found in group E. This occupies 
mangroves fringing the deserts of north-western Australia E22 ; mangroves, coastal 
forest and second growth from northern Australia to the Bismarcks and Solomons 
E23-26 (especially on very small islands) ; and second growth in the highlands of 
New Guinea E27. This specialization has allowed it to penetrate the ranges of the 
deep-forest forms P. schlegelii, P. soror and P. pectoralis H48. Although the two 
males E25 and H48 in the Bismarcks are similar in appearance, they differ markedly 
in habitat, conspicuousness and song (Dahl, 1899). As a consequence of their 
different habitat-preferences, members of group E must be exposed to different 
micro-climates from the neighbouring forest races. The special conditions in desert- 
fringing mangroves (presumably humid, yet subject to extremes of temperature) 
may explain the apparently contradictory changes of phenotype in north-western 
Australia E22 (p. 187). 

The variability of the hybrid populations on small islands off Shortland E26 may 
well be maintained by selection in relation to habitat differences. These populations 
are separated from one parental stock (C9) by less than three miles, from the other 
(dahli E25k) by almost two hundred. The latter occupies islets such as these, which 
elsewhere in the Solomons have no Pachycephala. Presumably the hybrids are better 
adapted to small islands than are the pure forms of group C. A surprising feature of 
the hybrid sample is that the smallest males have the whitest throats (like the small 
dahli) and the largest the yellowest throats (like the large Shortland race). In 
isolation, genetic linkage alone could keep the recombination classes scarce in the 
population for a few generations only. This would imply that at the time the known 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 191 

specimens were collected (1927) the hybrid populations were very young indeed, in 
which case they may by now have become partly stabilized. Supposing the situation 
to be of longer standing than this implies, the observed co- variation might be achieved 
by continued gene-flow from the parental stocks, or by selection in favour of the 
parental genotypes, or both. Recolonization by dahli must be almost unknown, 
while birds may arrive from Shortland rather frequently. In view of the failure of 
group C to colonize habitats such as are occupied by the hybrids, it seems probable 
that the dahli genotype is being maintained by selection, in the face of gene-flow 
from Shortland. 

Variation in bill size 

The absolute and relative length and stoutness of the bill varies greatly within 
the superspecies, to an extent which in many groups of birds would be considered to 
warrant generic separation. But the colour-patterns of the males are so clearly 
allied to one another that even Mathews (1930) places all these forms in a single genus. 
The variation shown by the bill is largely independent of the major discontinuities, 
and certain regularities apparent in a cursory study (pp. 186 and 188) show it to be at 
least partly adaptive. It is therefore relatively unimportant in the study of relation- 
ships within the group, and has not been dealt with in this paper. 

Except in long series, the finer geographical variation is to some extent obscured by 
individual variation and the inaccuracies of measurement (especially of bill depth) . 
The tables (p. 217) give some idea of the range in length and stoutness. Evidently 
very different bills (e.g. massive in Tenimber H42, slender in the Louisiades H47 and 
stubby in Tasmania F29) must be best adapted to taking correspondingly different 
food. But almost nothing is known about geographical differences in diet. The 
Guadalcanal race Cn takes considerably larger insects on the average than the 
smaller race, with a shorter and finer bill, on San Cristobal C17 (Cain & Galbraith, 
1956). Dahl (1899) records that of the two stocks in the Bismarcks, dahli E25 (with 
slightly the longer bill) takes a proportion of vegetable matter, while the race on the 
large islands H48 does not. 

CONCLUSIONS 

The very different plumage patterns of the sexes provide a large number of more 
or less independent characters which vary in stability and systematic importance, 
from the " qualitative " characters which unite males of the schlegelii assemblage 
to the slight differences in pigmentation and dimensions which distinguish closely- 
related populations on neighbouring islands. This makes it possible to study relation- 
ships within the superspecies despite the independent origin and loss of even the 
most stable characters, since the local co- variation of several relatively labile characters 
can be of equal importance. Most of the stable characters are provided by the male 
pattern, and most of the more plastic ones by that, of the female. The relationships 
suggested in this paper could not all have been arrived at by considering one sex 
alone, and it may not be possible to decide with any certainty the affinities of hen- 
feathered species such as P. simplex, P. sulfuriventer and P. philippensis. In 



i 9 2 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

organisms which do not show such diversity in conventional museum material, a 
genetic situation of equal complexity could only be interpreted by bringing in further 
characters (whether of internal anatomy, cytology, genetics, biochemistry, physiology, 
ecology or behaviour), which are most desirable in this group also, to test the validity 
of conclusions based entirely on a study of skins (cf. Wilson & Brown, 1953). 

The combination of great colonizing ability (implying dispersal) and divergence of 
neighbouring populations (implying philopatry) shown by the P. pectoralis super- 
species is remarkable, though not unprecedented. However, P. pectoralis is unique 
in having so many and so diverse forms, all of which are strictly allopatric, and between 
the most dissimilar of which there has been extensive gene-exchange. In this super- 
species hybridization and sympatry are largely independent of phylogenetic relation- 
ships, cutting across the division into schlegelii and soy or assemblages. It seems to 
be impossible to predict whether or not any given pair of representatives would 
interbreed on coming together, by considering only their resemblance and relation- 
ship. Purely local adaptations to the environment, and the manner of their meeting, 
are perhaps important in determining the outcome (p. 179). Although at the species 
level we have an objective criterion, not found higher or lower in the systematic 
hierarchy, for the relative status of any two populations which come in contact, 
it may not always be possible to use this criterion quite consistently in delimiting a 
given species. Strictly there are no biospecies, but only biospecific relations between 
sympatric populations. In the P. pectoralis superspecies, however, the three species 
P. schlegelii, P. soror and P. pectoralis can be satisfactorily delimited — although the 
different relationships between groups E and H in the Louisiades and Bismarcks 
(p. 168) suggest that in other groups it may be necessary to draw species-limits more 
arbitrarily. Morphological analogy is still the only available yardstick in determining 
the status of isolated representatives ; but the co-existence of closely-related forms 
in New Guinea, and interbreeding between more distant relatives in the archipelagos, 
demonstrate that other factors must be taken into account. Though at present we 
can only speculate on the nature of these factors, we need not consider as distinct 
species all those geographical representatives which are more different than the most 
similar pair of related sympatric species. It is justifiable and expedient to admit a 
wider range of representative forms to a single species than is the current practice. 
(In accordance with this consideration the Guadalcanal representative of Cichlomis 
whitneyi Mayr has been described as a subspecies (Cain & Galbraith, 1955), 
although the differences between it and the form in the New Hebrides are greater 
than those between many sympatric pairs of warblers.) 

The pattern of variation shown by the superspecies is interpreted as the result of 
colonizations by two stocks which diverged in New Guinea and attained biospecific 
relations with one another there, yet interbred freely in the surrounding archipelagos. 
The Lesser Sunda Isles, Moluccas and Solomons seem to have been colonized indepen- 
dently at an early date, from western and northern New Guinea, while the peculiar 
forms in northern Fiji and Samoa represent colonists from the Solomons (from 
which the New Caledonian race may also have received a contribution). Southern 
Melanesia is populated by forms which must have come from Australia at a later date 
(and which probably in turn colonized San Cristobal). Gaps in the range of these 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 193 

groups, from the Ban da Sea through the western Papuan islands and Santa Cruz 
to southern Fiji and Tonga, are occupied by a relatively undifferentiated stock which 
must have expanded relatively recently, and has formed hybrid populations with all 
the older groups with which it has come in contact. Finally, the stock which in 
northern Australia had become ecologically specialized for life in coastal and second- 
growth formations, has thereby been enabled to penetrate similar habitats in New 
Guinea and the eastern Papuan islands — sometimes merging completely with resident 
forms, sometimes remaining more or less isolated from them. 

This distributional history accords well with the avif aunal peculiarities of the sub- 
regions within Australasia, exemplifying several trends. For example, P. pectoralis 
is essentially an Australasian bird — but it slightly transgresses Wallace's Line and 
is stopped, not by the edge of the Sunda shelf, but presumably by the moister 
conditions and richer avifauna of western Java. Mayr (1944a) has shown that for 
the avifauna as a whole the Lombok Strait is merely the most effective single barrier 
in the series presented by water gaps and climatic differences along the Sunda Isles 
route. As is true of many birds, the species is represented by a peculiar form on the 
Tenimber Isles. The invasion from the Cape York peninsula of the relatively dry 
areas of southern and south-eastern New Guinea is paralleled by several species 
(e.g. Pachycephala rufiventris, Myiagra rubecula ; and see distribution maps in Mayr, 
19446). 

In the south-west Pacific the history of the species is representative for important 
elements of the avifauna. The Solomons are occupied by an ancient and peculiar 
endemic group, derived from New Guinea and showing extreme variation from island 
to island, whereas the Bismarcks have evidently been colonized only comparatively 
recently. A stock from Australia has occupied southern Melanesia and continued 
to San Cristobal (where the endemic form exemplifies several trends), and the New 
Hebrides are occupied by rather uniform populations. Both the northern Moluccas 
and the Solomons have well marked forms, belonging to the schlegelii assemblage and 
contrasting with those of the Banda Sea and eastern Papuan islands; but these 
are independently derived from P. schlegelii rather than directly related to one another. 
Faunal affinities between the two regions have been pointed out (Zeuner, 1943, 173 
for Troides and Hale Carpenter, 1953, 149 for Euploea (Lepidoptera) ; Voous & van 
Marie, 1949, for Coracina), and others might be suggested (e.g. Eos , sensu stricto , 
and " Eos " cardinalis ; Dicaeum erythrothorax and D. aeneum ; Rhipidura rufiventris 
cinerea-obiensis and Rh. cockerelli). The authors quoted postulate a continuous 
island chain, broken by the northwards drift of New Guinea as recently as the 
Pliocene ; but the explanation put forward for Pachycephala may be more acceptable — 
common origins in New Guinea, with parallel evolution under the effects of similar 
climates, avifaunas and degrees of isolation. 

No mention has been made of primitive characters. These can, of course, be recog- 
nized only from their occurrence and co- variation, not on a priori grounds. Clearly 
the immediate common ancestor of the superspecies was not hen-feathered but 
sexually dimorphic, and sexual dimorphism has been lost independently three times 
in P. pectoralis (and in the opposite way by P. flavifrons) . Other striking characters 
are similarly debarred from consideration by their scattered or peripheral distribution. 



194 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

It is reasonable to suppose that the pvoto-schlegelii showed the black chin and over- 
lapping throat-feathers which characterize members of the assemblage derived 
from it. But, without making unjustifiable assumptions about the possibility of 
independent origin or secondary loss of characters, it does not seem that anything 
can usefully be said about the other characters of this form, or about the common 
ancestor of both assemblages. Since the striking male variants from the standard 
pattern mostly reduce its distinctiveness, and since degenerative changes under relaxed 
selection probably proceed more quickly than selective enhancement of the pattern's 
visual properties, it is rather more likely that in any given instance the direction of 
evolution was away from the standard. But not even as much as this can be said of 
the labile female pattern. 

SUMMARY 

i. Pachycephala pectoralis, P. soror, P. schlegelii and P. flavifrons are considered 
to belong to a single superspecies, represented by a doublet in New Guinea (P. soror 
and P. schlegelii) with which P. pectoralis overlaps slightly. Standard patterns for 
males and females are pragmatically defined, and character-geographies of a number 
of the more clearly-defined variants given. Despite the independent origin and loss 
of characters, natural groups of subspecies can be distinguished (though not diagnosed) 
by considering the whole constellation of characters, weighted according to their 
co-variation. However, extensive secondary intergradation makes some of the 
boundaries between groups vague. 

2. A great range of forms is connected by intergradation, and must be considered as 
a single species (P. pectoralis) , although in New Guinea a pair of close relatives co-exist. 
It is concluded that conspecific allopatric forms may be more different than sympatric 
species, and P. caledonica of New Caledonia and P. melanops of Tonga are accordingly 
admitted to P. pectoralis (as is P. " orpheus " teysmanni of Salayer). The ability 
to interbreed is shown to be largely independent of relationship, and the species- 
limits to cut across the phylogenetic classification. 

3. Current criteria for the recognition of subspecies are criticized, and a higher 
standard of difference applied, in a subspecies-arrangement of P. pectoralis designed 
to reveal more clearly the uniformities and discontinuities. It is not suggested, 
however, that this arrangement should be adopted at present, and a list of sub- 
species to be recognized according to current practice is provided in the appendix. 

4. The distributional history of the superspecies is considered, and it is concluded 
that P. soror and P. schlegelii are descended from former geographical representatives 
within New Guinea. Near-standard subspecies of P. pectoralis in Australia, Southern 
Melanesia and elsewhere are closely related to P. soror, yet intergrade extensively 
with the peculiar endemic forms of the Lesser Sunda Isles, Moluccas, Solomons and 
Fiji — whose affinities are rather with P. schlegelii. But this view does not necessarily 
imply reticulate evolution, in the sense of a breakdown of established interspecific 
barriers. 

5. As might be expected from theoretical considerations and studies on geographical 
variation in other animals, few correlations with climate or habitat can be detected 
within the insular range of P. pectoralis. In Australia, however, regularities are 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 195 

apparent which agree with the well-established ecological rules of variation in 
homoiothermous animals ; and certain trends are apparent in forms which inhabit 
high altitudes, small islands, and islands where there are few bird species. There are 
in addition progressive character-changes in several insular areas, but most of these 
are interpreted as the result of secondary intergradation between forms of different 
origin. The geographical pattern of variation in these areas is consistent with the 
hypothesis of gene-flow, with peripherally-isolated forms shielded from its effects. 
It is possible that certain characters tend to appear especially in forms of hybrid 
origin. 

CHECKLIST 

Synonyms given by Mathews (1930) and Mayr (1932a, b, 1941a and 1954a) are 
not quoted here. A list of the subspecies to be recognized according to current 
practice is given on p. 205. 

PACHYCEPHALA SCHLEGELU Schlegel 

1 Pachycephala schlegelii schlegelii Schlegel. 

Pachycephala Schlegelii Schlegel, 1871 (from MS von Rosenberg). Tijdschr. ned. 
dierk. Ver. 4, p. 43 — l'interieur de la Nouvelle-Guinee [Arfak Mountains, according to 
Mayr, 1941a, 149]. 

Range : mountains of the Vogelkop, and Wandammen Mountains. 

2 Pachycephala schlegelii cyclopum Hartert 

Pachycephala schlegelii cyclopum Hartert, 1930. Novit. zool. 36, p. 54 — Cyclops 
Mountains. 

Range : Cyclops Mountains. 

3 Pachycephala schlegelii obscurior Hartert 

Pachycephala schlegelii obscurior Hartert, 1896. Novit zool. 3, p. 5 — Eafa District 
[Owen Stanley Mts.]. 

Pachycephala schlegelii viridipectus Hartert & Paludan, 1936. Mitt. zool. Mus. 
Berlin, 21, p. 203 — Kunupi [Weyland Mts.]. 

Range : (a) Weyland, Nassau and Oranje Mountains (" Snow Mountains ") 
intergrading with (b) Saruwaged and Sepik Mountains, and mountains of south- 
eastern New Guinea. 

PACHYCEPHALA SOROR Sclater 
1 Pachycephala soror soror Sclater 

Pachycephala soror Sclater, 1873. Proc. zool. Soc. Lond. p. 692 — Atam [Hatam], 
Arfak Mountains. 

Range : mountains of the Vogelkop. 
ZOOL. 4, 4. 14 



ig6 the pachycephala pectoralis superspecies 

2 Pachycephala soror klossi Ogilvie-Grant 

Pachycephala soror klossi Ogilvie-Grant, 1915. Ibis Jubilee Suppl. No. 2, p. 88 — 
the Utakwa Valley. 

Range : (a) mountains of northern New Guinea (Mamberano) and Weyland, 
Nassau and Oranje Mountains (" Snow Mountains "), intergrading with (b) Hagen, 
Bismarck and Saru waged Mountains. 

3 Pachycephala soror bartoni Ogilvie-Grant 

Pachycephala soror bartoni Ogilvie-Grant, 1915. Ibis Jubilee Suppl. No. 2, p. 89 — 
British New Guinea [Type : Owen Stanley Range, 5,000 ft.]. 

Range : mountains of south-eastern New Guinea and Goodenough Island. 



PACHYCEPHALA PECTORALIS (Latham) 
Ai Pachycephala pectoralis fulviventris Hartert 

Pachycephala fulviventris Hartert, 1896. Bull Brit. orn. CI. 5, p. 47 — Sumba. 
Range : Sumba. 

A2 Pachycephala pectoralis javana Hartert 

Pachycephala pectoralis javana Hartert, 1928. Bull. Brit. orn. CI. 48, p. 88 — Mt. 
Ardjuino, East Java. 

Range : (a) eastern Java, intergrading via (b) Bali with A3. 

A3 Pachycephala pectoralis fulvotincta Wallace 

Pachycephala fulvotincta Wallace, 1863. Proc. zool. Soc. Lond. p. 492 — Flores. 
Pachycephala pectoralis jubilarii Rensch, 1929. J. Orn. Lpz. Festschr. p. 202 — Alor. 

Range : (a) Sumbawa and Flores, (b) Lomblen, Pantar and Alor. 

A4 Pachycephala pectoralis everetti Hartert 

Pachycephala everetti Hartert, 1896. Novit. zool. 3, p. 170 — Insula Djampea. 
Pachy cephala pectoralis atrom.aculataM.eise, 1929. J. Orn. Lpz. 77, p. 448 — Kalao tua. 

Range : Djampea, Kalao tua and Madu. 

A5 Pachycephala pectoralis teysmanni Biittikofer 

Pachycephala teysmanni Biittikofer, 1893. Notes Leyden Mus. 15, p. 167 — Macassar, 
South Celebes [corrected to Salayer by Meyer & Wigglesworth, 1898, Birds of Celebes, 
2, p. 397]- 

Range : Salayer. 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 197 

B6 Pachycephala pectoralis mentalis Wallace 

Pachycephalia mentalis Wallace, 1863. Proc. zool. Soc. Lond. p. 30 — Bat j an et 
Gilolo [Type : Batchian]. 

Pachycephala pectoralis gilolonis Kuroda, 1938. Tori. 10, p. 114 — Halmahera. 

Range : Morotai, Halmahera and Bat j an. 

By Pachycephala pectoralis tidorensis van Bemmel 

Pachycephala pectoralis tidorensis van Bemmel, 1939. Treubia, 17, p. 99 — Tidore. 

Range : Tidore and Ternate. 

B8 Pachycephala pectoralis obiensis Salvadori 

Pachycephala obiensis Salvadori, 1878. Ann. Mus. Stor. nat. Genova, 12, p. 330 — in 
Obi. 

Range : Obi Islands. 

C9 Pachycephala pectoralis bougainvillei Mayr 

Pachycephala pectoralis bougainvillei Mayr, 1932. Amer. Mus. Novit. No. 522, p. 10 
— Bougainville Island, Solomon Islands. 

Range : Buka, Bougainville and Shortland. 

Cio Pachycephala pectoralis orioloides Pucheran 

Pachycephala orioloides Pucheran, 1853. Voy Pole Sud. Zool. 3, p. 57 — iles Salomon 
(San- Jorge). 

Pachycephala pectoralis pavuvu Mayr, 1932. Amer. Mus. Novit. No. 522, p. 15 — 
Banika Island, Pavuvu or Russel group, British Solomon Islands. 

Range : (a) Choiseul, (b) Ysabel, St. George and Florida Islands, (c) Russel 
Islands. 

Cn Pachycephala pectoralis cinnamomea (Ramsay) 

P.[seudorectes] cinnamomeum Ramsay, 1879. Nature, Lond., 20, p. 125 — Guadalcana. 
Range : Guadalcanal. 

C12 Pachycephala pectoralis sanfordi Mayr 

Pachycephala sanfordi Mayr, 1931. Amer. Mus. Novit. No. 504, p. 22 — Malaita 
Island, British Solomon Islands. 

Range : Malaita. 



C13 Pachycephala pectoralis melanonota Hartert 

Pachycephala melanonota Hartert, 1 
Lavella I., Central Group of the Solom 

Range : (a) Vella Lavella, (b) Ganonga. 



Pachycephala melanonota Hartert, 1908. Bull. Brit. orn. CI. 21, p. 106 — Vella 
Lavella I., Central Group of the Solomon Islands. 



i 9 8 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

C14 Pachycephala pectoralis melanoptera Mayr 

Pachycephala pectoralis melanoptera Mayr, 1932. Amer, Mus. Novit. No. 522, p. 18 
— Tetipari, central Solomon Islands. 

Range : Rendova and Tetipari. 

C15 Pachycephala pectoralis centralis Mayr 

Pachycephala pectoralis centralis Mayr, 1932. Amer. Mus. Novit. No. 522, p. 15 — 
Vangumi Island, central Solomon Islands. 

Range : Kulambangra, New Georgia, Vangunu and Gatukai. 

C16 Pachycephala pectoralis feminina Mayr 

Pachycephala feminina Mayr, 1931. Amer. Mus. Novit. No. 486, p. 25 — Rennell 
Island. 

Range : Rennell. 

C17 Pachycephala pectoralis christophori Tristram 

Pachycephalus christophori Tristram, 1879. Ibis, 4th ser., 3, p. 441 — Makira Harbour, 
San Cristoval, Solomon Islands. 

Range : (a) San Cristobal, (b) Santa Anna. 

D18 Pachycephala pectoralis graeffii Hartlaub 

Pachycephala grcBffii Hartlaub, 1866. Ibis, new ser., 2, p. 172 — Viti-levu. 
Pachycephala (?) optata Hartlaub, 1866. Ibis, new ser., 2, p. 172 — Ovalau. 

Range : (a) Viti Levu and Waia, intergrading through (b) Ovalau with D2ob. 

D19 Pachycephala pectoralis aurantiiventris Seebohm 

Pachycephala aurantiiventris Seebohm, 1891. Ibis, 6th ser., 3, p. 96 — Bua in 
Vanua Levu. 

Pachycephala pectoralis ambigua Mayr, 1932. Amer. Mus. Novit. No. 531, p. 16 — 
Rambi Island, Fiji Islands. 

Range : (a) Yanganga and most of Vanua Levu, intergrading via (b) Thaukan- 
drove Peninsula and (c) Kio and Rambi with D2oa. 

D20 Pachycephala pectoralis torquata Layard 

Pachycephala torquata Layard, 1875. ' Proc. zool. Soc. Lond. p. 150 — Taviuni. 
Pachycephala pectoralis koroana Mayr, 1932. Amer. Mus. Novit. No. 531, p. 15 — 
Koro Island, Fiji Islands. 

Range : (a) Taviuni (intermediate between D19C and D2ob) and (b) Koro. 



D21 Pachycephala pectoralis bella Mayr 

Pachycephala 
vara Island. 

Range : Vatu vara. 



Pachycephala pectoralis bella Mayr, 1932. Amer. Mus. Novit. No. 531, p. 14 — Vatu 
vara Island. 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 199 

E22 Pachycephala pectoralis melanura Gould 

Pachycephala melanura Gould, 1842. Proc. zool. Soc. Lond.,p. 134 — North coast of 
Australia [Derby, according to Mathews 1920, p. 229]. 

Eopsaltria hilli Campbell, 1910. Emu, 10, p. 168 — Hecla Island, Parry Harbour, 
North-West Australia. 

Pachycephala melanura bynoei Mathews, 1918. Aust. avian Rec. 3, p. 136 — Port 
Hedland. 

Range : (a) North West Cape to De Grey River, (b) Broome, (c) King Sound, 
(d) Hecla Island (off Cape Bougainville) and Napier Broome Bay — clinal series, 
confined to mangroves. 

E23 Pachycephala pectoralis violetae Mathews 

Pachycephala gutturalis violetae Mathews, 1912. Aust. avian Rec. 1, p. 76 — West 
Northern Territory [Daly R., according to Mathews 1920, p. 224]. 

Range : coasts and offshore islands of Arnhem Land and Gulf of Carpentaria, 
from Daly River to Normanton. 

E24 Pachycephala pectoralis spinicauda (Pucheran) 

Pteruthius spinicaudus Pucheran, 1853. Voy. Pole Sud, Zool. 3, p. 58 — l'ile 
Warriors [Torres Str.]. 

? Pachycephala salomonis Oustalet, 1877. Bull. Soc. philom. Paris, 6th ser., 12, p. 95 
— des iles Salomon [see p. 207]. 

Range : (? west coast of Cape York Peninsula), Cape York, islands in Torres 
Strait and coastal and second-growth formations in southern New Guinea from 
Merauke eastwards, probably intergrading with E25 near Hall Sound. 

E25 Pachycephala pectoralis dahli Reichenow 

? Pachycephala innominata Salvadori, 1881. Ornit. Pap. Mol. 2, p. 222 — in Papu- 
asia — ins. Teste {Ramsay). 

Pachycephala melanura dahli Reichenow, 1897. Orn. Mber. 5, p. 178 — Credner- 
Inseln, Raluan. 

Pachycephala pectoralis neuhausi Stresemann, 1934. O rn - Mber. 42, p. 24 — 
Sinabiet [Malie]. 

Pachycephala pectoralis fergussonis Mayr, 1936. Amer. Mus. Novit. No. 869, p. 2 
— Fergusson Island, D'Entrecasteaux Archipelago. 

Range : (a) ? south-eastern New Guinea, from Hall Sound to Milne Bay, (b) ? Teste 
Island, (c) Fergusson Island, (d) Long Island, (e) Witu Islands, (f) islands in Bungula 
Bay, New Britain, (g) Talele, Vatom, Duke of York and Credner (Palikuru) Islands, 
and shores of Blanche Bay, New Britain, (h) Nusa Island (off Kavieng, New Ireland), 
(j) Malie Island, Lihir group, (k) Nissan Island. 

E26 Pachycephala pectoralis whitneyi Hartert 

Pachycephala pectoralis whitneyi Hartert, 1929. Amer. Mus. Novit. No. 364, p. 14 
— Whitney Island [type designation and discussion attached in error to Pachycephala 
implicata] . 

Range : Whitney, Momalufu and Akiki Islands, east of Shortland — variable 
hybrid population between E25 and C9. Related populations, or pure populations 
of E25, may remain to be discovered elsewhere in the northern Solomons (see p. 208). 



200 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

E27 Pachycephala pectoralis balitn Rand 

Pachycephala pectoralis balim Rand, 1940. Amer. Mus. Novit. No. 1072, p. 8 — 
Balim River, altitude 1,600 meters ; Snow Mts., Netherland New Guinea. 

Range : second growth in the Balim and Bele Valleys, northern slopes of Mount 
Wilhelmina. 

F28 Pachycephala pectoralis fuliginosa Vigors & Horsfield 

[Pachycephala] fuliginosa Vigors & Horsfield, 1827. Trans Linn. Soc. Lond. 15, 
p. 241 — South coast of New Holland [Port Lincoln, according to Mathews, 1920, p. 
208]. 

Pachycephala occidentalis Ramsay, 1878. Proc. Linn. Soc. N.S.W. 2, p. 212 — 
Western Australia [Albany, according to Mathews, 1920, p. 209]. 

Range : (a) south-western Australia, west of a line through Geraldton, the Wongan 
Hills, Lake Grace and Esperance, (b) South Australia (Eyre and Fleurieu Peninsulas 
and Kangaroo Island), intergrading via (c) Victorian mallee with F3oa. 

F29 Pachycephala pectoralis glaucura Gould 

Pachycephala glaucura Gould, 1845. Birds of Australia, 2, part 18, p. 65 — Van 
Diemen's Land and the islands in Bass's Straits. 

Range : Tasmania (except the forests of the south-west) and islands in Bass 
Straits. 

F30 Pachycephala pectoralis pectoralis (Latham) 

Muscicapa pectoralis Latham, 1801. Index Orn. Suppl., p. 51 — Nova Hollandia 
[Port Jackson, according to Mathews 1920, p. 208]. 

Pachycephala gutturalis youngi Mathews, 191 2. Novit. zool. 18, p. 313 — Victoria 
[Lai Lai, according to Mathews 1920, p. 209]. 

Range : (a) Victoria east of a line from Heytsbury to Castlemaine, probably 
intergrading with (b) New South Wales. The range extends west of the Great 
Dividing Range into the Riverina district of southern New South Wales. Probably 
confined to eucalyptus forest, and riverine forest in savannah woodland. 

F31 Pachycephala pectoralis queenslandica Reichenow 

Pachycephala queenslandica Reichenow, 1899. Orn. Mber. 7, p. 8 — Nord Queens- 
land [Bellenden Kerr, according to Mathews, 1920, p. 209]. 

Pachycephala gutturalis ashbyi Mathews, 191 2. Novit. zool. 18, p. 313 — South 
Queensland [Blackall Ranges, according to Mathews, 1920, p. 209]. 

Range : (a) extreme north-eastern New South Wales (Richmond River) and 
southern Queensland (north to Mackay and Whitsunday Island), (b) Cairns district. 
Probably confined to rain forest. 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 201 

F32 Pachycephala pectoralis contempta Hartert 

Pachycephala contempta Hartert, 1898. Bull. B.O.C. 8, p. 15 — Lord Howe Island. 
Range : Lord Howe. 

F33 Pachycephala pectoralis xanthoprocta Gould 

Pachycephala xanthoprocta Gould, 1837. Proc. zool. Soc. Lond., p. 149 — in Nova 
Cambria Australi, apud oram orientalem [error for Norfolk Island according to Mathews, 
1928, Birds of Norfolk & Lord Howe Islands, p. 40]. 

Range : Norfolk Island. 

G34 Pachycephala [pectoralis] caledonica (Gmelin) — see p. 174. 

Muscicapa caledonica Gmelin, 1789. Syst. Nat. 1, p. 944 — nova Caledonia. 

Eopsaltria variegata Gray, 1859. Proc. zool. Soc. Lond. part 27, p. 162 — Island of 
Nu. 

Pachycephala morariensis Verreaux & des Murs, i860. Rev. Mag. Zool. p. 393 — 
[le] camp de Morari [New Caledonia]. 

Range : New Caledonia and Isle of Pines. 

G35 Pachycephala pectoralis littayei Layard 

Pachycephala Littayei Layard, 1878. Ann. Mag. nat. Hist. 5th ser., 1, p. 375 — 
Lifu, New Caledonia [!]. 

Range : Lifu and Uvea, Loyalty Islands. 

G36 Pachycephala pectoralis cucullata (Gray) 

Eopsaltria cucullata Gray, 1859. Cat. Birds Trop. Is. Pacific, p. 21 — New Hebrides 
(Aneitum) . 

Range : Aneitum. 

G37 Pachycephala pectoralis chlorura Gray 

Pachycephala chlorurus Gray, 1859. Cat. Birds Trop. Is. Pacific, p. 20 — New 
Hebrides (Erromango, Aneiteum) [restricted to Erromango by Mayr, 19326, p. 3]. 

Pachycephala intacta Sharpe, 1900. Ibis, 7th ser., 6, p. 343 — Sandwich Bay, Malli- 
collo. 

Pachycephala pectoralis brunneipectus Mayr, 1932. Amer. Mus. Novit. No. 531, p. 4 
— Epi Island. 

Pachycephala pectoralis banksiana Mayr, 1932. Amer. Mus. Novit. No. 531, p. 6 
— Vanua Lava, Banks Islands. 

Pachycephala pectoralis efatensis Mayr, 1938. Amer. Mus. Novit. No. 986, p. 2 — 
Efate Island, New Hebrides. 

Range : New Hebrides and Banks Islands, north of Tanna : (a) Erromango, (b) 
Efate and Nguna, (c) Mai, Tongariki, Epi, Lopevi, Pauuma and Ambrym, (d) 
Malekula, Malo, Espiritu Santo, and Dolphin Island, intergrading through (e) Omba, 
Raga and Maewo with (f) Gaua, Vanua Lava and Ureparapara. 



202 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

G38 Pachycephala pectoralis vanikorensis Oustalet 

P\achycephald\ vanikorensis Oustalet, 1877. Bull. Soc. philom. Paris, 6th ser., 
12, p. 95 — l'ile Vanikoro. 

Range : Vanikoro. 

H39 Pachycephala pectoralis calliope Bonaparte 

Pachycephala calliope Bonaparte, 1850. Conspic. Gen. Av. 1, p. 328 — Timor. 
Pachycephala melanura arthuri Hartert, 1906. Novit. zool. 13, p. 299 — Wetter. 

Range : (a) Timor and Samau, (b) Wetar. 

H40 Pachycephala pectoralis sharpei Meyer 

Pachycephala Sharpei Meyer, 1884. S.B. Isis Dresden, Jahr 1884, Abhandl., 
p. 36 — ins. Babbar. 

Range : Babar. 

H41 Pachycephala pectoralis dammeriana Hartert 

Pachycephala melanura dammeriana Hartert, 1900. Novit. zool. 7, p. 17 — Dammer 
Island. 

Range : Damar. 

H42 Pachycephala pectoralis fuscoflava Sclater 

Pachycephala fuscoflava Sclater, 1883. Proc. zool. Soc. Lond. p. 198 — Larat, ins. 
Tenimberensem. 

Range : Tenimber Islands. 

H43 Pachycephala pectoralis macrorhyncha Strickland 

Pachycephala macrorhyncha Strickland, 1849. Contr. Orn. (Jardine), p. 91 — 
Amboina. 

Pachycephala macrorhyncha alfurorum Stresemann, 1914. Novit. Zool. 21, p. 132 
— Gunung Sofia (Mittel-Seran) . 

Range : (a) Ceram, (b) Amboina. 

H44 Pachycephala pectoralis buruensis Hartert 

Pachycephala melanura buruensis Hartert, 1899. Bull. Brit. orn. CI. 8, p. 32 — Buru. 
Range : Buru. 



H45 Pachycephala pectoralis clio Wallace 

Pachycephala c< 
[restricted to the 

Range : Sula Islands 



Pachycephala clio Wallace, 1862. Proc. zool. Soc. Lond., p. 341 — Sula and Buru 
[restricted to the Sula Islands by Hartert, 1899. Bull. Brit. orn. CI. 8, p. 33]. 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 203 

H46 Pachycephala pectoralis pelengensis Neumann 

Pachycephala melanura pelengensis Neumann, 1941. Zool. Meded. 23, p. 112 — 
Peleng. 

Range : Peleng and Banggai Islands. 



H47 Pachycephala pectoralis collaris Ramsay 

Pachycephala collaris Ramsay, 1878. Proc. Linn. Soc. N.S.W. 3, p. 74 — Courtance 
Island, South-East coast, New Guinea ; torn, cit., p. 281 — Teste Island [see p. 206]. 

Pachycephala rosseliana Hartert, 1898. Bull. Brit. orn.Cl., 8, p. 8 — Rossel Island. 

Pachycephala pectoralis misimae Rothschild & Hartert, 191 8. Novit. zool. 25, 
p. 311 — St. Aignan or Misima Island. 

Range : (a) Conflict, Begum and Egum groups (and Coutance or Teste Islands?), 
intergrading through (b) Misima and the Deboyne group with (c) Rossel. 



H48 Pachycephala pectoralis citreogaster Ramsay 

? Saxicola merula Lesson, 1828. Voy. Coquille (Duperry) Zool. 1, pt. 2, p. 662 — 
la Nouvelle-Irlande, aux environs du Port Praslin [see p. 209]. 

Pachycephala citreogaster Ramsay, 1876. Proc. Linn. Soc. N.S.W. 1 p, 66 — New 
Britain and adjacent islands. 

Pachycephala pectoralis sexuvaria Rothschild & Hartert, 1924. Bull. Brit. orn.Cl. 
44, p. 50 — St. Matthias Island (Mussau). 

Range : (a) Umboi and New Britain, (b) New Ireland and Feni, (c) Lavongai, 
(d) Mussau. 



H49 Pachycephala pectoralis ottomeyeri Stresemann 

Pachycephala pectoralis ottomeyeri Stresemann, 1933. Orn. Mber. 41, p. 116 — 
Komat auf Lihir. 

Range : Lihir Island. 



H50 Pachycephala pectoralis tabarensis Mayr 

Pachycephala pectoralis tabarensis Mayr, 1955. Amer. Mus. Novit., No. 1707, 
p. 35 — Tabar Island, Tabar group. 

Range : Tabar Island. 



H51 Pachycephala pectoralis goodsoni Rothschild & Hartert 

Pachycep 
p. 296 — Ma 

Range : Manus 



Pachycephala pectoralis goodsoni Rothschild & Hartert, 1914. Novit. zool. 21, 
p. 296 — Manus. 



204 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

H52 Pachycephala pectoralis ornata Mayr 

Pachycephala pectoralis ornata Mayr, 1932. Amer. Mus. Novit. No. 531, p. 8 — 
Santa Cruz [Ndeni], Santa Cruz Islands. 

[Pachycephala] atrata Mayr, 1932. Amer. Mus. Novit. No. 531, p. 10 — nomen nudum 
[used by Mayr in MS.]. 

Range : (a) Ndeni, (b) Reef, Duff and Swallow groups. 

H53 Pachycephala pectoralis utupuae Mayr 

Pachycephala pectoralis utupuae Mayr, 1932. Amer. Mus. Novit. No. 531, p. 8 — 
Utupua, Santa Cruz Islands. 

Range : Utupua. 

H54 Pachycephala pectoralis kandavensis Ramsay 

Pachycephala kandavensis Ramsay, 1876. Proc. Linn. Soc. N.S.W. 1, p. 65 — 
" Kandavu ". 

Range : Kandavu group and Mbengha. 

H55 Pachycephala pectoralis vitiensis Gray 

Pachycephala vitiensis Gray, 1859. Cat. Birds Trop. Is. Pacific, p. 20 — Feejee 
Islands (Island of Ngau). 

Range : Ngau. 

H56 Pachycephala pectoralis lauana Mayr 

Pachycephala pectoralis lauana Mayr, 1932. Amer. Mus. Novit. No. 531, p. 12 — 
Ongea Levu Island, Lau Archipelago, Fiji Islands. 

Range : Ongea Levu, Fulanga and Wangava, southern Lau Archipelago. 

H57 Pachycephala pectoralis melanops (Pucheran) 

Eopsaltria melanops Pucheran, 1853. Voy. Pole Sud., Zool. 3, p. 56 — Vavao. 
Range : Vavau group and Late, Tonga. 

PACHYCEPHALA FLAVIFRONS (Peale) 

Eopsaltria flavifrons Peale, 1848. U.S. Explor. Exped. Birds (subsequently with- 
drawn), p. 96 — Upolu. 

Range : Upolu and Savai, Samoa. 

Subspecies to be recognised according to current usage 

Subspecies which are distinguished from one another by measurements alone, and 
which are near the borderline of subspecific distinctness under the seventy-five 
percent rule, are bracketed together and the junior name indicated by an asterisk. 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 



205 



Pachycephala schlegelii : 

1 schlegelii 

2 cyclopum 

3 obscurior 

Pachycephala soror : 

1 soror 

2 klossi 

3 bartoni 

Pachycephala pectoralis : 
Ai fulviventris 
A2 javana 

{A3 a fulvotincta 
A3b jubilarii* 
A4 everetti 
A5 teysmanni 
B6 mentalis 
B7 tidorensis 
B8 obiensis 
C9 bougainvillei 
Cioa-b orioloides 
Cioc pavuvu 
Cn cinnamomea 
C12 sanfordi 
C13 melanonota 
C14 melanoptera 
C15 centralis 
C16 feminina 
C17 christophori 
Di8a graeffii 
Di8b optata 
D19 aurantiiventris 
D2oa torquata 
D2ob koroana 
D21 fo//a 
E22a bynoei 
E22b-c melanura 
E22d A2 - //i 
E23 violetae 
E24 spinicauda 



{E25C fergussonis* 
E25CI-J 6?aA/i 
E26 whitneyi 
E27 balim 
F28a occidentalis 
F28b-c fuliginosa 
F29 glaucura 
F3oa youngi 
F3ob pectoralis 
F3ia ashbyi 
F3ib queenslandica 
F32 contempta 
F33 xanthoprocta 
G34 caledonica 
G35 littayei 
G36 cucullata 
G37a chlorura 
G37b-f intacta 
G38 vanikorensis 
H39a calliope 
H39b arthuri 
H40 sharpei 
H41 dammeriana 
VL/±2 fuscoflava 

{H43a alfurorum* 
H43b macrorhyncha 
H44 buruensis 
H45 cJio 
H46 pelengensis 
H47a-b collaris 
H47C rosseliana 
H48a-c citreogaster 
H48CI sexuvaria 
H49 ottomeyeri 
H50 tabarensis 
H51 goodsoni 
H52 ornata 
H53 utupuae 
H54 kandavensis 
H55 vitiensis 
H56 lauana 
H57 melanops 



NOTES 

Pachycephala schlegelii viridipectus Hartert & Paludan (3b) 

The differences between viridipectus and obscurior 3a are too slight for subspecific 
separation (cf. Mayr, m &#.), besides which they intergrade smoothly (Mayr & 
Gilliard, 1954). 



206 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

Pachycephala pectoralis jubilarii Rensch (A3D) 

From a study of A.M.N. H. material, Mayr (in litt.) concludes that the size difference 
between jubilarii and fulvotincta A3a is sufficient for subspecific separation. 

Pachycephala pectoralis atromaculata Meise (= A4) 

Mayr (in litt.) finds no difference in colour between atromaculata and everetti, and 
my few measurements suggest no size difference adequate for separation. 

Pachycephala pectoralis gilolonis Kuroda (= B6). 

Mayr (in litt.) finds no difference between gilolonis and mentalis. 

Pachycephala pectoralis ambigua Mayr (Digc) 

The undesirability of recognizing two variable hybrid subspecies (ambigua and 
torquata D2oa) between aurantiiventris Dioa and koroana D2ob has been discussed 
(p. 176). Since ambigua intergrades smoothly with aurantiiventris it seems best to 
combine them, although the end populations are very distinct. 

Female of bynoei (E22a) 

There are four females from Cossack in the White Collection of the National 
Museum of Victoria. Although no comparative material of melanura E22C was 
available, these appeared to agree well with the female of that form. The desirability 
of a comparative description has been pointed out to the Museum authorities. 

Inter gradation of spinicauda (£24) with dahli (£25) 

A single female from Dalena, Hall Sound (A.M.N. H. Reg. No. 329999) differs from 
typical spinicauda females, and approaches those of dahli, in having the underparts 
much yellower and less ochraceous (cf. Rand, 1940). 

The subspecies on Teste Island 

Ramsay (18780, 74) described Pachycephala collaris (H47a) from Cou(r)tance 
Island, off the coast of south-eastern New Guinea. Later (1878b, 281) he recorded 
P. melanura ( = spinicauda E24) on Coutance, and gave the locality of collaris 
(without further comment) as Teste Island, off the extreme south-eastern tip of New 
Guinea. He further described from Teste a form which Salvadori later (1881, 222) 
named P. innominata (? E25b) from this description. The single specimen is described 
as having an ashy-grey tail and slaty-black occiput, and the yellow collar " inter- 
cepted on the head and neck ". 

The type of collaris and the female described by Ramsay are in the British Museum 
(Natural History), Reg. Nos. 95.12.24.2 & 4 respectively. Neither bears a field 
label, but both are reputedly from Coutance. Two males (B.M.(N.H.) Nos. 78. 10. 19.5 
& 6) bear field labels giving their locality as Teste, and No 6 is recorded as collected 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 207 

by G. W. Baiston Ingham, one of the collectors mentioned by Ramsay (18786, 241). 
These specimens agree well with males of spinicanda, fergussonis and dahli E24 & 
25c-k. 

There are no further specimens of P. pectoralis from Teste or Coutance Islands 
in the B.M.(N.H.), the A.M.N.H., or the Australian Museum. It seems most 
probable that both are inhabited by black- tailed forms of group E., and that the 
lost type of innominata was a specimen moulting into adult plumage. Conceivably 
Ramsay's notorious unreliability over localities extends in this instance to the 
description, and " ashy-grey " should refer to the wings. P. innominata Salvadori 
must be considered unidentifiable. The type locality of collaris remains to be 
determined. For the present it seems best to accept the evidence of the original 
description, and the label of the type, giving the locality as Coutance. 



Pachycephala pectoralis fergussonis Mayr (E25C) 

The two known specimens of fergussonis are distinctly larger than typical dahli, 
and slightly deeper yellow beneath. However, dahli from Long Island E25d (and Teste 
E25b ?) approach them in size. In view of the geographical variation in size within 
dahli, and the slightness of the colour difference, it is best to submerge fergussonis 
in dahli until females are available for comparison (cf. Mayr, in Hit). 



Pachycephala salomonis Oustalet. (? = E24) 

The locality of the single (male) specimen was given (Oustalet 1877, 95) as the 
Solomon Islands, although d'Urville's " Voyage au Pole Sud " in the " Astrolabe " 
and " Zelee " did not stop there. From an examination of the type by Professor 
Berlioz, Mayr (1932a, 21) concluded that salomonis is a synonym of dahli 
Reichenow, 1897 (E25),but that (owing to the unreliability of the locality and the het- 
erogynism of closely related races) it must be considered unidentifiable. Comparison 
of the type with males of dahli, spinicauda and citreogaster, kindly undertaken by 
Professor Berlioz, confirms that it is inseparable from those of the violetae-spinicauda- 
dahli aggregate E23-25, and quite different from citreogaster H48. Mayr (1955, 34, 
and in litt.) supposes the type to have been collected at Port Praslin, New Ireland, 
and the name therefore to be a synonym of citreogaster. This error springs from 
that (Mayr, 1932a) of supposing that the type was collected on D'Urville's earlier 
" Voyage de V Astrolabe" , which called at Port Praslin. 

Even if the locality of salomonis were known, the principle of conservation (Copen- 
hagen Decisions, 1953, 25) would debar the use of the name, since it seems not to 
have been used since its publication. From the itinerary of the voyage it seems most 
probable that the type was collected at Port Essington (violetae E23) or the Torres 
Strait (spinicauda E24) , where the vessels actually called ; though it might have been 
brought by canoe from Teste Island or Nissan (dahli E25), or even from somewhere 
in the northern Solomons (cf. whitneyi E26 ?). In the circumstances, P. salomonis 
must be considered unidentifiable. 



208 THE PACHYCEPHALA PECTORALIS SUPERSPECIES 

White-throated Pachycephala in the Solomons 

Besides the hybrid race whilneyi E26 on small islands west of Shortland, there are 
indications that other populations related to dahli E25 may remain to be discovered 
in the northern Solomons. Hartert (1926, 46) records a specimen from Munia, south- 
west of Fauro in the Bougainville Strait. A male (B.M.(N.H.) Reg. No. 36.4.20.14) 
collected on or near Buka (Moyne- Chaplin, 22nd December, 1935) agrees well with 
dahli and with white-throated males of whitneyi, except that it has a small black 
chin-spot. 

Pachycephala pectoralis brunneipectus, banksiana and efatensis Mayr (G37C, 
f &b) 

Though separable in long series of females from different islands, all the populations 
from Efate to the Banks Islands G37b-f should be combined in intacta Sharpe (cf . 
Mayr, in litt.). 

Pachycephala macrorhyncha alfurorum Stresemann (H43a) 

The size difference between alfurorum and macrorhyncha K^b seems to be sufficient 
for subspecific separation (Mayr, in litt.), as my measurements tend to confirm. 

Pachycephala collaris Ramsay (1147a) 

In view of the uncertainty about the type locality of collaris (p. 206), Professor 
Mayr has suggested that a redescription of the original specimens may be useful. 

The following descriptions must be read in conjunction with those of the standard 
patterns (p. 138). Colours are cited according to the code of Villalobos-Dominguez 
& Villalobos (1947). The type was compared with two males of misimae 1147b 
from Misima (B.M.(N.H.) Nos. 99.5.20.6 & 7) and one of rosseliana H47C (1917.11. 
21. 1) ; Ramsay's female with two of misimae from Misima (99.5.17.43 and 
99.5.20.5). Unfortunately, they were not critically compared with a male of 
collaris from East Island, nor a female of rosseliana, both borrowed from the American 
Museum of Natural History. 

Adult male. Type (B.M.(N.H.) Reg. No. 1895. 12. 24. 2). 

Throat-feathers with little or no grey at bases (as misimae and rosseliana). 

Underparts about OOY/OY.17.12 : misimae similar, rosseliana a little more 
golden, towards OOY. 

Underside of tail a little more olivaceous, less fuscous, than in rosseliana, misimae 
intermediate. 

Collar somewhat washed with brownish-olive on hind-neck : in rosseliana, much 
narrower and quite olivaceous on hind-neck, misimae intermediate. 

Mid-back about YYO . 5 . 12 : rosseliana darker and greener, about YYO/Y . 3 . 12 , 
misimae paler and slightly greener, about YYO(Y) .7. 12 . 

Edges of remiges (worn) greyer, edges of upper wing-coverts yellower, centres of 
wing-feathers paler than in rosseliana — misimae agrees with collaris. 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 209 

Tail (only 5 rectrices remain) yellowish olive, central rectrices without black ; 
remainder with vague brown patches (somewhat broken up into transverse bars) 
subterminally on inner webs ; shafts brown, paling basally. In misimae central 
rectrices have vague brownish barring ; remainder have blackish brown patches 
occupying most of the inner webs ; shafts darker brown. In rosseliana, central 
rectrices are dark olive with long blackish patches along the shafts on the inner 
webs ; remainder are brownish black with dull olive edges (widest towards the bases 
of the outer webs) ; shafts black, becoming dark brown basally, 

In rosseliana only, the rump, upper tail-coverts and tail are somewhat washed with 
brown. 

Adult female. (95 . 12 . 24 . 4) . 

Throat white with brownish fringing (as in misimae) . 

Gorget narrow, pale and vaguely-defined, vinous grey in colour (about SO.16.2 ) 
as in 99 . 5 . 20 . 5 of misimae ; in 99 . 5 . 17 . 43 it is wider, deeper, more sharply defined 
beneath and browner (about OOS . 12 . 4 ) . 

Underparts rich yellow, about OY.17.12 on mid-belly, with lower breast and 
flanks somewhat olivaceous to brownish ; 99 . 5 . 17 . 43 agrees ; 99 . 5 . 20 . 5 is much 
paler, about OY . 18 . io° on mid-belly. 

Upperparts very brown, as in 99.5.17.43 (crown about OOS.6.5 , mid-back 
about 0.4. 12 ) ; 99 . 5 * 20 . 5 is much less brown, with crown greyer (about OOS .5.3°) 
and mantle greener (about YYO.4.10 ). 

Edges of wing-feathers slightly greyer, less rufous, than in misimae. 

Tail intermediate between 99 . 5 . 17 . 43 (browner) and 99 . 5 . 20 . 5 (greener). 

Pachycephala pectoralis misimae Rothschild & Hartert (1147b) 

Although it shows signs of gene-flow from rosseliana H47C, this form is nearest to 
collaris H47a, with which it should be combined (cf. Mayr, in litt.). 

Saxicola merula Lesson (? = H48b) 

The locality of the type, a juvenile, was given by Lesson as Port Praslin, on the 
south-east coast of New Ireland. This would make the name the senior synonym 
of citreogaster , though the principle of conservation would require its suppression. 
But many specimens brought back by French expeditions of the late eighteenth and 
early nineteenth centuries are wrongly localized, and several species recorded from 
Port Praslin do not in fact occur in the Bismarcks (Mayr, in litt.). Salvadori (1881, 
219) questioned the locality, pointing out the resemblance of the type to juveniles 
of macrorhyncha (Amboina H43b). Juveniles of citreogaster and macrorhyncha 
probably cannot be separated with certainty, and Mayr, who has examined the type, 
considers it to be unidentifiable. 

ACKNOWLEDGEMENTS 

With great pleasure I record my indebtedness to the following : 

Dr. A. J. Cain introduced me to the principles and methods of systematics, suggested 
the study of Pachycephala pectoralis and the research on geographical variation of 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 



which it forms a part, and has been a constant source of advice. Professor Ernst 
Mayr has greatly encouraged me in the study of a group on which he is the authority, 
and liberally made me free of his knowledge and experience : my indebtedness to his 
published and unpublished work will be evident throughout the text of this paper. 
Dr. Dean Amadon has been to great trouble to answer a list of queries from the 
material in the American Museum of Natural History. Professor Jean Berlioz has 
been most helpful in comparing the type of Pachycephala salomonis, and discussing 
the nomenclatural problem. Dr. Allen Keast has unsuccessfully sought the type 
of Pachycephala innominata in the Australian Museum. Dr. Cain, Professor Mayr, 
Mr. R. E. Moreau and Dr. H. W. Parker have read drafts of this paper at various 
stages, and made invaluable criticisms and suggestions. Mr. H. O. Ricketts has 
carefully checked the final draft, detecting many lapses and obscurities, and helped 
me in proof-reading. Mr. R. J. Drumm, O.B.E., has greatly helped in seeing the 
paper through the press. Both the British Museum (Natural History) and the 
American Museum of Natural History have allowed me to borrow many specimens. 
The paper was prepared between 1952 and 1955, at the Department of Zoology 
and Comparative Anatomy, University of Oxford, under Professor A. C. Hardy, 
F.R.S. Working facilities were also granted at the Edward Grey Institute of Field 
Ornithology by Dr. David Lack, F.R.S. , and at the Bird Room of the British Museum 
(Natural History) by Mr. J. D. Macdonald. Professor Hardy and the Department 
of Scientific and Industrial Research gave financial assistance during the preparation 
of the paper, and field work in the Solomons was financially supported by the Percy 
Sladen Trustees and the Parliamentary Grants Committee of the Royal Society. 

MEASUREMENTS 

In addition to my own measurements, Professor Mayr has most kindly put at my 
disposal most of those taken by himself (for Mayr, 19316, c; 1932a, b; 1936, 1938, 
1941a, 1944c and 1955) and by Mrs. Kate Jennings (for Mayr, 1954a). Several 
specimens were measured both by Mayr or Jennings and by myself, while I measured 
many others twice. The distributions of the discrepancies (between Mayr's and 
Jennings' and my own first measurements on the one hand, and my definitive 
series on the other) indicate to what extent these results are comparable or 
repeat able. 

Wing : self 2 — Mayr mean — 0-07 mm., s.d. ±1-20 mm 
self 2 — Jennings ,, — 0-18 ,, ,, ±0-95 



(27 measurements) 

(14 » ) 

(148 „ ) 

(26 

(15 
(148 

the mean discrepancies 



self 2 — selfi ,, + 0-40 ,, ,, i 0*24 

Tail: self 2 — Mayr „ +0-25 „ „ ±0-98 

self 2 — Jennings ,, + 0-08 ,, ,, ±2-11 

self 2 — selfj ,, — 0-41 „ ,, ±0-51 

Although several individual discrepancies exceed 2 mm. 
are small. However, several of the standard deviations differ to a high degree 
of significance (by the variance-ratio test), and it seems unwise to combine Mayr's 
and Jennings' series with my own. I have inserted them at the appropriate points 
in the table, prefixed by M or J respectively. Asterisks indicate measurements in 
my series which appear also in Mayr's or Jennings'. Since weight determinations are 
not subject to the same personal bias, I have combined our series for Ci7a. Weights 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 211 

for Cn were taken by Cain and myself, the remainder by the Whitney Expedition 
of the American Museum of Natural History. 

Series of five measurements or less are given in full, with replications indicated 
in parenthesis — e.g. 76, 78-5, 80(2), 81-5. Longer series are summarized in the form 
mean ^ standard deviation [number in series) — e.g. 86-8 ± 2- 10 (13). 

Recognizably distinct populations are distinguished in the tables by their ciphers, 
and by subspecies names (not all of which are admitted on p. 205) where appropriate. 
Weights and wing and tail lengths are given separately for adult males and females. 
The sexing of Mayr's specimens, those of the O.U.(D.Z.) Solomons Expedition 
(Cn & 17a), and the hen-feathered series C16 and F33 have been accepted for this 
purpose. However, much of the remaining material is not sexed, and some determina- 
tions are questionable. Therefore only specimens showing feathers belonging to the 
adult male plumage have been taken as adult males ; and only those sexed as females 
and showing female plumage, without male or juvenile feathers, as adult females. 

In the few forms of which I had adequate series of males, females and juveniles, 
reliably sexed, there was little indication of significant and constant differences 
between sex and age groups, in the measurements of bills and tarsi (values for Ci7a 
are analysed on p. 218). I have therefore risked the introduction of some bias, 
in order to have longer series, by combining the measurements of all specimens 
except nestlings. Mayr's raw data contain very few individual measurements of 
bills and none of tarsi, and I have not repeated the indications of size range pub- 
lished in the papers cited. 

Weight in grams 

My measurements taken to the nearest 0-5 gm., with a long-scale spring balance 
(Gibb balance), calibrated in the field. 





Males 


Females 


Cioa orioloides 


49'9 ± 2-62 (17) 


42, 44. 5o 


Cn cinnamomea . 


5i-5 ± 2 -24(i9) 


44 -9±i-8i(8) 


C12 sanfordi 


51-3 ± 2-25 (18) 


49*5 ± 2 -oo (19) 


Ci7a christophori . 


33'5 ± 2-96(45) 


32-1 ±2-36 (20) 


Ci7b 


34. 35 (2) 


— 


E25k dahli. 


29-0 ± 1-69 (8) 


27, 28, 29 



Wing and tail lengths, in millimetres (pp. 212-216) 

My wing lengths taken to nearest 0-5 mm., from wing-bend to tip of longest 
primary (of left wing wherever possible), with wing pressed flat. 

My tail lengths taken to nearest 0-5 mm., from tip of longer central quill to 
insertion of central quills in common sheath. 

Tarsus and culmen lengths, and bill depth, in millimetres (pp. 217-219) 

My tarsus lengths taken to nearest 0-5 mm., along outer side of tarsus, from groove 
of intertarsal joint to eminence near plantar angle of hind toe. 

My culmen lengths taken to nearest 0-5 mm., chordwise, from tip to angle of 
culmen with skull. 

My bill depths taken to nearest 0-5 mm., perpendicular to tomium at hind edge 
of nostril, from culmen to lower edge of mandible (bill fully closed) . 

zool. 4, 4. 15 



212 



THE PACHYCEPHALA PECTORALIS SUPERSPECIES 



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Australasian archipelago. Trans, zool. Soc. Lond. 25 : 107-184. 




h Fig. 6) showing original colonizations by the 
nblage (black) and the soror assemblage (stippled). Group H of P. 
1 (see Fig. 6, inset right). 





E27 ^"---,^24 



X 



(53 



Dl« DM H57 



■ '••.»^i.> C9 C|0 CII H52 H53 

B8. ll!^/:H"V?^">0 CMr^ : 5' // '. V\G38X;. C 

(CH^ V^^^^^^----------' \^ N HH H \ 5 

^ H H 




&, 



fi'm 




Diagram (for comparison with Fig. 6) showing areas of secondary inter- 
ion between subspecies-groups of P. pectoralis (stippled), and minor gene-flow 



Sketch-map of the distribution of the superspecies. Solid lines, boundaries of P. schlegelii, P. soroy and P. fiavifrons ; broken lines, 
' i of the subspecies-groups of P. pectoralis; dotted lines, boundaries of subspecies (omitted where continental subspecies intergrade). 
Inset left, distribution of P. soror and subspecies-group E of P. pectoralis in the Papuan region. Inset right, relative positions of the subspecies- 
groups of P. pectoralis : groups A to G stippled, widespread group H black. 




2 7 NOV 






A REVISION OF THE 

LAKE VICTORIA HAPLOCHROMIS 

SPECIES (PISCES, CICHLIDAE) 

PARTI: H. OBLIQUIDENS HILGEND., 
H. NIGRICANS (BLGR.), 

H.NUCHISQUAMULATUS (HILGEND.) 
AND H. LIVIDUS, SP. N. 



P. H. GREENWOOD 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 4 No. 5 

LONDON: 1956 



A REVISION OF THE LAKE VICTORIA 
HAPLOCHROMIS SPECIES (PISCES, CICHLIDAE) 

PART I: H. OBLIQUIDENS HILGEND., 
H. NIGRICANS (BLGR.), 

H. NUCHISQUAMULATUS (HILGEND.) 
AND H. LIVIDUS, SP. N. 



BY 

P. H. GREENWOOD 

East African Fisheries Research Organization, Jinja, Uganda. 



Pp. 223-244 ; 2 Text-figs. 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 4 No. 5 

LONDON: 1956' 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 4, No. 50/ the Zoological series. 




PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued November, 1956 Price Six Shillings 



A REVISION OF THE LAKE VICTORIA 
HAPLOCHROMIS SPECIES (PISCES, CICHLIDAE) 

PART I: H. OBLIQUIDENS HILGEND., 

H. NIGRICANS (BLGR.), 

H. NUCHISQUAMULATUS (HILGEND.) 

AND H. LIVIDUS, SP. N. 

By P. H. GREENWOOD. 

CONTENTS 

Page 
Introduction ............. 226 

Haplochromis obliquidens Hilgend. . . . . . . . . .226 

Synonymy and description . . . ... . . . . .227 

Distribution ............ 229 

Ecology ............. 229 

Affinities and taxonomic status of the species . . . . . .230 

Study material and distribution records . . . . . . .232 

Haplochromis lividus sp. nov. .......... 232 

Synonymy and description . . . . . . . . .232 

Distribution . . . . . . . . . . . . 234 

Ecology 234 

Diagnosis ............. 236 

Study material and distribution records . . . . . . .237 

Haplochromis nigricans (Blgr.) . . . . . . . . . .237 

Synonymy and description . . . . . . . . .237 

Distribution ............ 239 

Ecology ............. 239 

Diagnosis ............. 240 

Study material and distribution records ....... 240 

Haplochromis nuchisquamulatus (Hilgend.) . . . . . . .241 

Synonymy and description . . . . . . . . .241 

Distribution ............ 242 

Ecology 242 

Diagnosis ............. 242 

Study material and distribution records ....... 243 

Summary .............. 243 

Acknowledgments ............ 243 

References .............. 243 

zool. 4, 5. 16 



226 REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 



INTRODUCTION 

The species described in this paper form a well-defined ecological group within the 
Haplochromis species flock of Lake Victoria. All feed principally by grazing on 
epiphytic and epilithic algae. 

As a group and severally they show obvious morphological adaptations to this 
particular feeding habit. Adaptation is most clearly seen in tooth-form and arrange- 
ment, which depart from those common to the majority of Haplochromis species. 

Several other ecologico-morphological groups have evolved in Lake Victoria. 
Their existence raises questions regarding the possibility of providing a realistic 
basis for subdividing the present phylogenetically amorphous arrangement of the 
species. 

There are, however, certain difficulties inherent in this procedure. A strictly 
morphological approach to sub-division is unworkable. Intergradation, rather than 
discreteness, of morphological group-characters might be said to typify this species 
flock. Such a situation is, however, not unexpected in a large group of oligophyletic 
origin which has undergone intense adaptive radiation during a short period of 
geological time (Regan, 1922 ; Greenwood, 1951). 

Also, although some morphologically distinct species-complexes occupy equally 
distinctive ecological niches, there are other morphologically-homogeneous groups 
which cut across any attempted ecological classification. 

Furthermore, in any ecologically-defined group there are grades of anatomical 
specialization such that the most and least specialized species are only with difficulty 
in eluded in a supra-specific category defined by morphological criteria alone. The 
species described here typify this situation. Tooth-form in H. obliquidens is unlike 
that of most species at present included in the genus Haplochromis. Yet three algal- 
grazing species are known, which partially bridge this morphological gap. At the 
opposite extreme H. nuchisquamulatus exhibits incipient dental adaptation only 
slightly removed from a generalized Haplochromis type. 

In Lake Victoria, then, there exist several nascent supra-specific groups which are 
more readily identified by ecological than morphological criteria. Since conventional 
taxonomic characters are, so to speak, also nascent, formal recognition of these 
categories is impossible. I propose, therefore, to recognize their biological and evolu- 
tionary significance only by drawing attention to their existence. 



Haplochromis obliquidens Hilgendorf, 1888 

Chromis {Haplochromis) obliquidens Hilgendorf, 1888, 5. B. Ges. naturf. Fr. Berlin, 76. 

Ctenochromis obliquidens, Pfeffer, 1897, Arch. f. Naturg., 63, 60. 

Tilapia obliquidens, Boulenger, 1898, Trans, zool. Soc, Lond., 15, 5. 

Hemitilapia bayoni Boulenger, 1908, Ann. Mus. Genova (3) 4, 6; Idem, 191 1, Ibid. (3) 5, 69; 

Idem, 1915, Cat. Afr. Fish., 3, 491, fig. 340. 
Haplochromis nuchisquamulatus (part), Boulenger, 1915, op. cit., 290. 
Clinodon bayoni (Blgr.), Regan, 1920, Ann. Mag. nat. Hist. (9), 5, 33. 
Haplochromis obliquidens (part), Regan, 1922, Proc. zool. Soc, Lond. 188. 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 227 

The holotype of Haplochromis obliquidens could not be examined ; it is amongst 
those specimens, once housed in the Berlin Museum, and which cannot be located 
at present. However, the characters noted in Hilgendorf s original description are 
diagnostic. 

Through the courtesy of Dr. D. Guiglia (Museo Civico di Storia Naturale, Genoa) 
I was able to study the holotype of Hemitilapia bayoni Boulenger, and thus to confirm 
Regan's synonymy of this species with H. obliquidens. 

On the other hand, I cannot agree with Regan's tentative synonomy of Hemitilapia 
materfamilias Pellegrin, 1913, and Haplochromis obliquidens (Regan, 1922). Re- 
examination of H. materfamilias type specimen revealed that Pellegrin's original 
description is misleading, particularly in respect of the dentition, and that the species 
should be referred to Macropleurodus bicolor (Blgr.) (Greenwood 1956). 

Description. Based on fifty-seven fishes (size range 48-89 mm. standard length) 
including the holotype of Hemitilapia bayoni and five specimens in the British 
Museum (Natural History). Three other British Museum (Nat. Hist.) specimens 
were examined, but are not included in the morphometric data. 

Since no marked allometry with size was determined for any character examined, 
measurements are given for the collection as a whole, with the exception of the smallest 
specimen, which is treated separately. 

Depth of body 33-4-41-2, mean (M) 37-5, length of head 29-4-34-0 (M = 32-3) 
per cent of standard length. Dorsal profile of head and snout straight ; fairly 
steeply sloping in most fishes but decurved in a few individuals. Preorbital depth 
12-5-17-4 (M = 15-2) per cent of head length ; least interorbital width 27-8-34-7 
(M = 31-8) ; snout as broad as or somewhat broader than long, rarely longer than 
broad, its length 26-6-33-3 (M = 29-2) per cent head length. Eye 29-1-33-3 (M = 
31-4) ; depth of cheek 19-0-25-0 ( M= 21-5) per cent head length. 

Caudal peduncle about ij times as long as deep ; 13-2-16-4 (M == 15-0) per cent 
of standard length. 

Corresponding ratios for the smallest individual (48 mm. S. L.) — not included in 
the mean values given above — are : Head 32-3 ; preorbital n-i ; interorbital 27-8 ; 
snout 27*8 ; eye 27-8 ; cheek 16-7 ; and caudal peduncle 16-6 per cent. 

Mouth short and horizontal or very slightly oblique ; posterior maxillary tip 
extending to the vertical from the anterior orbital margin, or almost so. Jaws equal 
anteriorly, the lower 31-6-41-6 (M = 37-2) per cent of head length ; its length/ 
breadth ratio from 1-1-1-7 (mode 1-4). 

Gill rakers short ; 8 or 9, rarely 7 or 10, on the lower limb of the anterior arch. 

Scales ctenoid ; lateral line interrupted, with 30 (f.6), 31 (f.31) or 32 (f.18) 
scales. Cheek with 3 (rarely 2 or 4) series of imbricating scales. 5 or 6 scales 
between the dorsal fin origin and the lateral line ; 5-7 between pectoral and pelvic 
fin insertions. 

Fins. Dorsal with 24 (f.25), 25 (f. 31) or 26 (f.i) rays, anal 10 (f.i), n (f.n), 12 
(f.43) or 13 (f.2), comprising XIV-XVI, 8-10 and III, 7-10 spines and soft rays for 
the fins respectively. First pelvic ray slightly produced and variable in its posterior 
extension, usually reaching the spinous anal fin in adults and occasionally to the soft 
part in ripe males. Caudal sub-truncate. 

zool. 4, 5. i6§ 



228 REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 

Teeth. Teeth forming the outer series are movably implanted and have slender 
necks with undivided, expanded, compressed and obliquely truncate crowns (Text- 
fig, i). In many specimens a few postero-lateral teeth in both jaws are bicuspid 
but otherwise retain almost the same crown form as the more anterior teeth ; in 
some the second cusp is incipient, but in others it is clearly differentiated. There is 
no correlation between length of fish and the presence or number of undifferentiated 
postero-lateral teeth. 




1 mm 



A weak positive correlation exists between the number of teeth in the outer series 
of the upper jaw, and standard length. 

S.L. (mm.) . . 48 55-64 . 65-74 • 75-87 

Tooth number . . 50 42-60 . 50-70 . 58-70 

Mean . . . .50 -54 -59 .64 

N .... 1 . 20 . 21 . 12 

The inner teeth are mostly tricuspid and arranged in 2-4 rows, with a distinct 
interspace separating them from the outer series. Some obtusely tricuspid teeth, 
and others differing only in their smaller size from those in the outer series, frequently 
occur in the first inner row. These obliquely truncate teeth are larger than their 
tricuspid associates. 

Alizarin preparations of two larvae (10 and 11 mm. total length) obtained from 
the mouth of a brooding female, show larval dentition to be comparable with that 
of H. macrops (Blgr.) H. prodromus Trewavas and Macropleurodus bicolor at an 
equivalent developmental stage. The teeth of H. obliquidens larvae differ 
considerably from the adult condition, being slender and setiform, with slightly 
recurved unicuspid crowns ; 14-16 outer teeth, aggregated medially, are present 
in the upper jaw. 

In certain fishes from Kisumu, it was noticed that the crowns of all teeth were 
coarse and irregular, and that their typical golden-brown coloration was replaced by 
black. Similar structural differences and discoloration have been observed in 
specimens of H. michaeli Trewavas from various localities. The cause of this 
aberrancy is unknown. 

Lower pharyngeal bone sub-equilaterally triangular, its dentigerous surface 
broader than long. The numerous teeth are fine, compressed and directed 
posteriorly, and have truncated crowns ; a small anterior cusp is present in all. 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 229 

Skeleton. Differs in no important respect from that of generalized Haplochromis 
species. Vertebrae : 14 + 17, 13 -f- 17, 13 + 16 or 12 + 17. 

Coloration in life : Breeding males. Ground colour bright yellow-green, becom- 
ing yellower ventrally ; chest and branchiostegal membrane blackish ; lips slightly 
irridescent. Dorsal fin yellow-grey, lappets red ; orange-red spots and streaks 
on the posterior spinous and entire soft parts. Anal with a pinkish flush ; 3 or 
4 yellow ocelli. Pelvics with black outer and clear or faint pink inner half. Non- 
breeding adult males are similarly coloured except that the body is more nearly 
olivaceous and the chest not darkened. Females and juveniles of both sexes. Ground 
colour silvery-yellow. Dorsal and caudal fins neutral ; anal and pelvic fins pale 
yellow. Darker, almost olivaceous females are known. 

Transverse banding occurs in both sexes, but is rarely apparent in life. 

Preserved material : Adult males. Dusky, the vertical bars partly or completely 
obscured. Dorsal fin sooty, lappets black, the posterior spinous and entire soft 
part maculate ; anal colourless ; pelvics dark laterally, pale mesially ; caudal 
maculate. Females and juveniles. Grey to brown, with or without six to ten narrow 
transverse bars on the flanks ; less frequently a faint mid-lateral stripe and a fainter 
stripe approximately following the upper lateral line. Fins colourless and immaculate. 

Distribution. Haplochromis obliquidens has been collected from many localities 
in Lake Victoria. It is also known from the Victoria Nile. 

Recently, Miss R. H. Lowe obtained a small sample of H. obliquidens from Lake 
Bunyoni (Uganda). Earlier reports (Worthington, 1932) indicated the probability 
that no Haplochromis were then present in this lake. It is presumed that those now 
occurring there were accidentally introduced on occasions when the lake has been 
stocked with Tilapia species ; that one of the two species now recorded is probably 
H. nigripinnis Regan (otherwise endemic to Lake Edward) and the other is H. 
obliquidens supports this assumption, since Tilapia have been introduced from both 
Lakes Edward and Victoria. 

The nine Lake Bunyoni H. obliquidens (size range 63-85 mm. S.L.) differ slightly 
from the Lake Victoria population in the following characters : body more slender, 
30-2-34-5 (M = 32-4) per cent of standard length ; the preserved coloration of 
sexually mature males is apparently more melanic ; in three specimens the outer 
series of teeth is entirely composed of bicuspids similar to the undifferentiated 
postero-lateral teeth of Lake Victoria fishes. In all other observed morphological 
characters the two populations are identical. 

Ecology : Habitat. Shallow littoral zone, particularly in the vicinity of emergent 
vegetation ; less commonly in the water-lily zone, over exposed sandy beaches 
and at the margin of papyrus swamps. There are indications, both from fishing 
and direct observation, that H. obliquidens may frequent rocky shore-lines, where 
the substrate is largely composed of broken rocks and boulders. The species has 
often been collected and seen around rock foundations of piers. 

Food. The intestine of H. obliquidens is long and much coiled (2J-3 times S.L.) ; 
stomach large and distensible. Stomach and intestinal contents of fifty-three 
individuals (size range 48-89 mm. S.L.) from various localities, have been examined. 

Diatoms comprised the main digested contents in forty-four individuals ; the 



230 REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 

genera principally recorded were : Melosira, Suririella, Gomphonema, Rhopalodia, 
Navicula and Cyclotella. 

Small fragments of plant epidermis occurred in the stomachs of twenty-nine fishes. 
The quantity ingested by individuals varied considerably. It was observed that, 
unless ruptured, most epidermal cells were apparently undigested. 

Blue-green algae, especially Rivularia and Microcystis, and less frequently 
Anabaena and Oscillatoria, were recorded from nineteen stomachs ; none of these 
plants showed signs of digestion. 

Filamentous green algae, chiefly Spirogyra and to a lesser extent Oedogoniutn, 
occurred in sixteen stomachs. No digestion was noted. 

The stomach contents of one individual comprised only partly digested fragments 
of Ephemeroptera larvae, probably taken at the time of their emergence. 
Fragmentary remains of both adult and larval insects were found in the intestines 
of three other fishes. 

The frequent occurrence of epiphytic algae and epidermal fragments of phanero- 
gams suggests that H. obliquidens feeds partly by scraping the surface of submerged 
leaves and stems. This supposition is confirmed by observations made on the 
feeding behaviour of these fishes in the lake ; the peculiar dentition of H. obliquidens 
would seem to be highly adapted for such habits. 

On the other hand, sand grains and bottom debris were also found in many 
stomachs ; indeed, it was often difficult to determine whether ingested plant frag- 
ments were the partly digested remains of epidermis scraped from living plants or 
whether they were derived from the semi-decayed debris which accumulates near 
dense plant stands. Probably H. obliquidens feeds both by grazing on plants and 
by utilizing plant material contained in the bottom detritus. In either eventuality 
it is clear that diatoms are the principal food organisms utilized, and that much 
ingested plant material is voided undigested. 

Although rather infrequent, the occurrence of insects in the pabulum could 
indicate that the species is partly facultative in its feeding habits and may utilize 
temporary and seasonal abundances of animal food. 

Breeding. Breeding behaviour and spawning sites of H. obliquidens are unknown. 
However, females carrying young in the buccal cavity have been obtained from 
most localities. 

The smallest sexually active fish was a female 61 mm. long ; above 68 mm. 
S.L., most individuals were found to be mature. 

Affinities and taxonomic status of the species 

Particular interest attaches to H. obliquidens, since although it is the type species 
of the genus its dental morphology is unique amongst the very numerous species of 
Haplochromis. Throughout this discussion the generic diagnosis is taken to be that 
prepared by Regan (1920) in which particular emphasis was laid on neurocranial 
osteology. Subsequently this definition has been modified by the recognition of 
several related genera distinguished from Haplochromis by their divergent dentition 
(Regan, 1922 ; Trewavas, 1938 ; Greenwood, 1956). 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 231 

Within the genus thus denned two types of outer teeth predominate, a unicuspid, 
conical form and a bicuspid compressed type. The common dental pattern is a 
single outer series distinctly separated from the inner series, usually comprising two 
or three rows anteriorly and a single row postero-laterally. 

Tooth form and pattern have played an important part in species discrimination 
and in the actual or attempted delimitation of supra-specific groups amongst Lake 
Victoria species. Extreme dental specialization, associated with osteological 
changes, characterises four of the five monotypic cichlid genera in this lake (Regan, 
1922 ; Greenwood, 1956), whilst less obvious dental characters were used in an 
attempt to subdivide the endemic Haplochromis into five genera (Regan, 1920) . Two 
years later, Regan abandoned this concept, reducing some of his genera to subgeneric 
rank and discarding others (idem, 1922). 

Because the dental morphology of H. obliquidens does not conform with that 
usual for Haplochromis, there might appear to be grounds, as Regan suggested 
(op. cit.), for recognizing at least one sub-genus to accommodate those species with 
unequally bicuspid or conical outer teeth. The sub-genus Ctenochromis (Pfeffer, 
1893) would be available for such species (Regan, loc. cit.). In that paper Regan 
first indicated H. astatodon Regan of Lake Kivu as providing a dental type, which 
although invariably bicuspid, linked the " obliquidens " tooth form with that of the 
commonly occurring Ctenochromis type. The teeth of H. astatodon exhibit some 
diversity in the degree to which they approach the " obliquidens " condition, but 
the greatest number of individuals has teeth approximating more closely to this 
type than to " Ctenochromis ". The common tooth form in H. astatodon may be 
likened to a bicuspid variant of typical H. obliquidens teeth ; indeed, similar teeth 
frequently occur postero-laterally in both jaws of H. obliquidens. 

Two other annectent species have since been found : H. annectidens Trewavas 
from Lake Nabugabo and a new species (described below) from Lake Victoria. 
Intra-specific variation in the tooth form of this latter species is as great as that of 
H. astatodon, but most individuals possess teeth similar to the undifferentiated 
postero-lateral teeth of H. obliquidens. 

It is clear, then, that although the teeth of H. obliquidens may represent an extreme 
form, intermediates linking them to the usual bicuspid Haplochromis type are found 
as the characteristic dentition in three extant species. The gap separating the most 
" obliquidens "-like teeth of H. astatodon and the new species from those of H. 
obliquidens is relatively slight ; it represents no more than the loss of a small cusp 
from an expansive, compressed and obliquely truncate crown. Less modified 
crown structure as seen in some teeth of these two species, grades through 
the condition found in H. nuchisquamulatus, into the more usual, acutely bicuspid 
form. 

Thus, the case for recognizing at least two sub-genera of Haplochromis on the 
basis of dental morphology (Regan, 1920 and 1922) is weakened. As was 
mentioned earlier, several ecologically defined groups, each comprising apparently 
related species, are known from Lake Victoria. In every case, the group shows 
certain morphological divergence from the generalized Haplochromis type, but no 
clear-cut gap has evolved which would allow for its formal recognition as a sub-genus. 



232 REVISION OF LAKE VICTORIA 

Study material and distribution records 

Museum and Reg. No. 

Genoa Museum. Holotype of Hemitilapia bayoni 
British Museum (N.H.) 1908, 10. 19.6 (paratype 

of H. bayoni) .... 
British Museum (N.H.) 1911, 3.3.80 
British Museum (N.H.) 1913, 9.30. 13-18 
British Museum (N.H.) 1956, 7.9.1-16 

,, ,, 17-20 

„ „ „ 21-27 

„ „ „ 28 

„ „ „ 29-32 

», M ,, 33-45 

„ „ „ 4^55 

„ „ „ 56-57 

,, ,, I69-I70 



HAPLOCHROMIS SPECIES 



Locality. 
Sesse Islands 

Sesse Islands 

Jinja (Ripon Falls) 

Lake Victoria 

Jinja (Pier) 

Beach near Nasu Point 

(Buvuma Channel) 
Grant Bay (Buvuma 

Channel) 
Napoleon Gulf, near 

Bugungu (opp. Jinja) 
Entebbe Harbour 
Kisumu, Kavirondo Gulf 
Mwanza, Capri Bay 
Godziba Island 
Kalagala, Victoria Nile 



Collector. 
Bay on. 

Bayon. 
Bay on. 
Bayon. 
E.A.F.R.O. 



Haplochromis lividus sp. nov. 

Haplochromis nuchisquamulatus (part), Blgr., 1915, Cat. Afr. Fish., 3, 290, Fig. 197. 
Haplochromis desfontainesii (part), Blgr., 1915, op. cit., 302. 
Haplochromis nubilus (part), Regan, 1922, Proc. zool. Soc, London, 164. 

Type specimen. A male go -f- 21 mm. from Bugungu (near Jinja), Uganda. 

Description. Based on seventy-seven fishes (size range 46-90 mm. S.L.) from 
Lake Victoria. Five specimens from Lake Kyoga are considered separately. 

Within the size range of individuals studied no character showed marked allometry 
with standard length or length of head ; measurements are therefore given for the 
whole collection with the exception of the smallest fish, which was not included 
when determining means. 

Depth of body 33-3-41-2 (M = 36-5) ; length of head 31-0-35-0 (M = 32-7) 
per cent of standard length. Dorsal head profile straight and moderately steeply 
sloping (ca. 45 °), rarely somewhat curved. Preorbital depth 12-0-16-7 (M = 14-7) 
per cent head length ; least interorbital width 26-2-33-3 (M = 29-7) ; snout as 
broad as long, its length 26-0-32-0 (M = 28-8) per cent of head. Eye 28-0-36-0 
(M = 31-4) ; depth of cheek 17-0-24-1 (M = 20-1) per cent head length. 

Caudal peduncle 1-1-1-7 (M = 1-4) times as long as deep ; 12-2-18-5 (M = 15-5) 
per cent standard length. 

Corresponding ratios for the smallest fish (46 mm. S.L.) are : Depth 39-0, head 39-0 
per cent of standard length. Preorbital 12-8, interorbital 23-2, snout 27-8 and 
cheek 16-7 per cent of head-length. Caudal peduncle 15-2 per cent of S.L. 

Mouth horizontal or slightly oblique ; posterior maxillary tip reaching the vertical 
to the anterior orbital margin or nearly so, and to the eye in some. Lips slightly 
thickened. Lower jaw 33-3-41*0 (M = 37-2) per cent of head, its length /breadth 
ratio 1-3-2-0 (mode i-6). 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 



233 



Gill rakers short, 8 or 9 (less frequently 7 or 10) on the lower part of the first arch. 

Scales ctenoid, lateral-line interrupted, with 30 (f.7), 31 (f.16), 32 (f.48), 33 
(f.5), or 34 (f.i) scales. Cheek with 2 or 3 (rarely 4) series. 5 or 6 scales between 
dorsal fin origin and the lateral line ; 5 or 6 between pectoral and pelvic fin insertions. 

Fins. Dorsal with 23 (f.i), 24 (f.34), 25 (f.40) or 26 (f.2) ; anal n (f.18), 12 (f.55), 
and 13 (f.4), rays, comprising XV-XVI 8-10 and III, 8-10 spinous and soft rays for 
the fins respectively. First pelvic ray produced, variable in its posterior extension, 
but reaching the spinous anal in most adults. Pectoral fins as long as, or slightly 
shorter than the head. Caudal sub-truncate. 

Teeth. In the form and pattern of its teeth, H. lividus departs from the generality 
of Haplochromis species. The anterior and antero-lateral teeth in the outer series 
are movably implanted and have slender necks (somewhat stouter than in H. 
obliquidens) with compressed, expanded and obliquely truncated, unequally bicuspid 
crowns. The posterior cusp shows some variation in size, but it is always smaller 
than the anterior, from which it is narrowly separated (Text-fig. 2). It should be 




1mm 





Fig. 2 



noted that these teeth bear a striking resemblance to the undifferentiated postero- 
lateral teeth of H. obliquidens. Posterolateral teeth in H. lividus are either similar 
to the anterior teeth or indistinguishable from the generalized acutely bicuspid 
type (Text-fig. 2G). Less frequently, unicuspid teeth occur in this position. A 
weak positive correlation exists between the number of teeth in the outer series of 
the upper jaw and standard length. 



S.L. (mm.) 
Tooth number 
Mean 


46 
36 
36 


• 56-65 
. 42-58 
50 


66-75 
38-66 

54 


76-85 

• 44-75 
61 


86-91 

52-66 

57 


N 


1 


8 


27 


33 


5 



234 REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 

Teeth forming the inner rows are invariably tricuspid and small. No obtusely- 
cuspidate teeth, or teeth similar to those of the outer series, have been observed 
(cf. H. obliquidens in which such teeth are frequently encountered). 

There is considerable variation in the number and disposition of inner rows. 
From 2-5 and from 2-4 series occur in the upper and lower jaws ; individuals with 
more than three rows usually have the interspace between outer and inner teeth 
greatly reduced or even absent, particularly in the upper jaw. 

Lower pharyngeal bone sub-equilaterally triangular ; dentigerous surface slightly 
broader than long. Pharyngeal teeth similar to those described for H. obliquidens. 

Osteology. That of a typical generalized Haplochromis species ; vertebrae 
14 + 16 (in two specimens). 

Coloration. Breeding colours of male H. lividus are perhaps the most distinctive 
morphological characteristic of the species and are not repeated or even approached 
in any other Lake Victoria Haplochromis. In preserved material their brilliance is 
lost. Sexually active males. Ground colour light olive-green shading to slate-grey 
ventrally ; flanks (including the dorsal aspects and in some, the nape) with a golden- 
red flush extending from the head to the caudal peduncle origin. Inter-orbital 
region of the head, the snout, lips and preorbital with a vivid, almost fluorescent, 
blue sheen, traces of which often extend onto the otherwise slate-grey lower jaw, 
lower preopercular limbs and the branchiostegal membrane. As far as can be 
determined, the intensity of this peculiarly intense head coloration is little 
influenced by the fishes' emotional state. On the other hand, its greatest extension 
is apparently manifest only in breeding fishes. 

Dorsal fin grey to sooty, slight indications of fluorescent blue can be detected in 
some individuals ; red streaks on the posterior spinous and entire soft part ; lappets 
orange-red. Caudal dark, with ill-defined red maculae concentrated proximally 
on the upper half. Anal dark, with 2-4 yellow ocelli. Pelvics black, becoming 
lighter on the medial third. Coloration of immature males is similar except that the 
blue head colour is less concentrated and intense, or it may even be absent. The 
flanks are also less intensely red. Females. Ground colour light grey-green, becom- 
ing silver ventrally. Dorsal, caudal and pectoral fins colourless or faintly yellow- 
grey. Anal and pelvic fins yellow. 

Preserved material : Males. Ground colour variable, usually grey. Dorsal 
caudal and anal fins clear or dark, the two former maculate as in life ; pelvics black. 
The blue head coloration is lost, but in most individuals it is faintly represented by 
a dead-white or ashen colour (at least in formalin fixed material preserved in spirit 
for five years). Females and immature males. Ground colour as above. All fins 
clear. From 5-7 transverse bars on the flanks ; the posterior pair rarely extend 
below the level of the lower lateral-line and are often joined by a short longitudinal 
stripe. Faint indications of a mid-lateral stripe are present in some individuals. Band- 
ing and striping are sometimes apparent in living fishes, but are intensified after death. 

Distribution. H. lividus is known from several localities in Lake Victoria. 
(See below.) 

Ecology : Habitat. Shallow littoral zone, especially in the vicinity of emergent 
and submerged vegetation, less frequently in the water-lily zone and at the margin 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 235 

of papyrus swamps ; the species is commonly encountered over rock foundations of 
piers. Thus, the habitat preferences of H. lividus are similar to those of H. 
obliquidens with which species it is usually captured. There are, however, indica- 
tions that H. lividus may inhabit the deeper littoral zone where H. obliquidens are 
relatively scarce. 

Food. The intestine is long (2-2 \ times standard length) and much coiled ; the 
stomach large and distensible. The stomach contents of sixty- two individuals 
(size range 56-90 mm. S.L.) from most localities have been examined. In general 
the food of H. lividus is similar to that of H. obliquidens. 

Diatoms of the genera Melosira and Rhopalodia comprised the predominating 
digested contents in the stomachs of forty-five fishes, and were significant in twelve 
others. 

Fragments of plant epidermis were found in thirty-three stomachs ; as in H. 
obliquidens the amount and fragment-size showed considerable variation. 

Filamentous green-algae, represented by Spirogyra, were recorded from only 
four fishes ; in none was there any indication of digestion. Blue-green algae 
(especially Rivularia and Microcystis) were found in twenty-four stomachs. Again, 
the algae were apparently not digested. 

Very fragmentary animal remains (Ostracoda, Crustacea [Decapoda] and Insecta 
[larval Chironomidae]) were recorded from sixteen individuals. 

The occurrence of Insecta (winged Hymenoptera) as the main stomach contents 
in twelve fishes collected contemporaneously at one station, is of particular interest. 
Besides insect remains, diatoms were well represented in these stomachs. This 
observation suggests that H. lividus may feed facultatively on animal food at times 
of local abundance. Insects also comprised the main contents of two other 
specimens, both from different localities. 

Feeding habits of H. lividus are probably similar to those of H. obliquidens. 
Direct observation shows the species to be a grazer on submerged plants and stones, 
whilst the occurrence of sand-grains and bottom debris in some stomach contents 
indicates occasional benthic feeding. 

Breeding. Breeding habits and sites are imperfectly known. Three females 
carrying young in the buccal cavity have been collected ; one from an exposed 
beach flanked by dense emergent vegetation and two from an off-shore water-lily 
stand. From one of these fishes twenty-five larvae of 12 mm. total length were 
recovered ; the other females had jettisoned the greater part of their broods. 

Affinities. Disregarding for the moment its peculiar male breeding coloration, 
H. lividus shows marked affinity with H. astatodon, H. annectidens and H. obliquidens, 
especially with regard to dental characteristics. Save H. annectidens, for which 
no information is available, the food of these species is similar, and composed mainly 
of epiphytic algae and plant debris (Poll and Damas, 1939, for food of H. astatodon). 

In fact, H. lividus, H. astatodon and H. annectidens seem to provide examples of 
herbivorous intermediates linking generalized and usually insectivorous Haplochromis 
species with the specialized algal-grazer, H. obliquidens. 

Within the species flock of Lake Victoria H. lividus shows some morphological 
relationship with H. nuchisquamulatus. Anatomical and dental characteristics 



236 REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 

might entitle H. nuchisquamulatus to consideration as the extant representative of an 
annectant form between H. lividus and the generalized species typified in Lake 
Victoria by H. nubilus (Blgr.) and H. macrops (Blgr.). 

The present distribution of species having " lividus "-like teeth requires little 
comment. Haplochromis astatodon is endemic to Lake Kivu, whose Haplochromis 
species flock has long been recognized as having well-defined Victorian affinities 
(Regan, 1921), although the possibility of convergent evolution in the two lakes 
cannot be entirely discounted. Haplochromis annectidens is endemic to Lake 
Nabugabo and is part of a small species group which could only have been derived 
from that of Lake Victoria (Trewavas, 1933). 

The presence within Lake Victoria of both H. lividus and its morphological 
derivative H. obliquidens is suggestive of an ancestor-descendant relationship. 
Before accepting this apparent phylogeny, due regard must be paid to the unique 
male breeding coloration of H. lividus. Baerends and Baerends van Roon (1950) 
expressed the opinion that male coloration plays an important part in species 
recognition amongst cichlids. Thus, we may assume the importance of male 
coloration as a barrier to interspecific mating. Field observations on the Haplo- 
chromis of Lake Victoria lend weight to this hypothesis. Although male colours and 
colour-patterns are broadly repeated in several species, no instance has yet been 
recorded of related species with identical or near identical male coloration breeding 
in the same habitat. 

Therefore, although the distinctive coloration of H. lividus might be used in 
argument against close relationship with H. obliquidens, it might equally well be 
interpreted as resulting from selection strengthening mating barriers between 
species which occupy similar habitats, especially if the species are closely related 
and of recent origin. 

Diagnosis. Haplochromis lividus differs from other Haplochromis in Lake 
Victoria in having distally compressed and expanded teeth whose crowns are 
unequally bicuspid and obliquely truncated. Dentition serves to distinguish this 
species from the fluviatile Haplochromis of East Africa. In life male coloration is 
the most obvious diagnostic character. 

From species with similar dental morphology H. lividus may be differentiated as 
follows : from H. astatodon by its larger eye /cheek ratio ; from H. annectidens by 
its slightly wider interorbital region and somewhat stouter, shorter teeth. In life 
coloration distinguishes H. lividus and H. astatodon ; live colours are unknown for 
H. annectidens. 

Five specimens from Lake Kyoga (Tilapia nubila B.M. (N.H.) reg. nos. 1911 .3.3. 
141-145 ; 60-66 mm. S.L.) have teeth and dental patterns of the H. lividus type, 
but differ from Lake Victoria specimens in the following characters : dorsal head 
profile steeper ; body deeper ; and greater depth of cheek (23-3-25-2, mean 24-1 
per cent head length). Should further collections from Lake Kyoga show that 
these fishes have H. lividus coloration (as is suggested in the preserved material) 
and should they also maintain the observed differences in morphology, then it will 
be necessary to recognize a distinct sub-species in that lake. 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 



237 



Study material and distribution records 



Museum and reg. 


no. 


Locality. 


Collector. 


British Museum 


(N.H.) 1906 


.5.30.318-320 


Entebbe 


Degen. 


British Museum 


(N.H.) 1956. 


7.9. 63-65 . 
„ ,, 58-62 . 


Jinja Pier 
Beach near Jinja 


E.A.F.R.O 


>> >> 


,, >, 


„ „ 66-74 • 


Napoleon Gulf, near 






(Type and paratypes) 


Bugungu (opp. Jinja) 




,, ,, 


>> »> 


» » 75-83 • 


Kirenia (near Jinja) 




,, ,, 


>> >> 


„ „ 84-94 • 


Entebbe Harbour 




" " 


" " 


„ „ 95-99 • 
,, ,, 100-102 


Beach near Nasu Point, 

Buvuma Channel 
Hannington Bay (Uganda) 




>> >> 


tt >> 


„ „ 103-124 


Grant Bay (Uganda) 




>> » > 


>> >> 


„ „ 125 


Mwanza, Capri Bay 




" 


,, 


,, ,, 126-128 


Majita (Tanganyika 
Territory) 





Haplochromis nigricans (Blgr.) 1906 

Tilapia nigricans (part) Blgr., 1906, Ann. Mag. nat. Hist. (7) 17, 448 ; Idem, 1907, Fish. Nile, 
518 ; Idem, 191 1, Ann. Mus. Genova (3) 5, 75 ; Idem, 1915, Cat. Afr. Fish., 3, 241, fig. 160. 
Tilapia simotes Blgr., 191 1, Ann. Mus. Genova (3) 5, 75 ; Idem., 1915, op. cit., 242, fig. 161. 
Neochromis nigricans (Blgr.), Regan, 1920, Ann. Mag. nat. Hist. (9) 5, 33. 
Haplochromis (Neochromis) nigricans (Blgr.), Regan, 1922, Proc. zool. Soc, London, 163. 

As Regan (1922) first showed, Boulenger's figure of Tilapia nigricans is misleading. 
It was prepared from a specimen distorted in preservation and consequently the 
head profile differs considerably from that of T. simotes. However, in the important 
characters of dental pattern and morphology both species are identical. If the head 
of the figured specimen is restored to its natural position the characteristically 
decurved profile of Haplochromis nigricans is apparent. 

Description. Based on fourty-four specimens (size range 49-94 mm. standard 
length) including holotypes of T. nigricans and T. simotes. Other specimens in the 
British Museum (Nat. Hist.) collections were examined but are not included in the 
morphometric data. The paratype of T. nigricans is clearly not referable to this 
species and should probably be placed in H. lividus. Its small size permits only 
tentative identification. Both skeletons in the British Museum (Nat. Hist.) are of 
H. nigricans. 

Depth of body 34-5-40-0 (mean 36-9), length of head 28-0-33-3 (M = 31-2) per 
cent of standard length. Dorsal head profile strongly decurved. Preorbital depth 
11-8-16-7 (M = 14-6) per cent of head length ; least interorbital width 25-0-31-5 
(M = 28-8) ; snout broader than long in most specimens of more than 65 mm. 
S.L., and as long as broad in smaller fishes, its length 26-3-35-2 (M = 30-4) per cent 
of head. Eye 25-9-33-3 (M = 30-0), depth of cheek 19-4-27-3 (M = 32-4) per cent 
of head. 

Caudal peduncle from i-i-i-8 (mode 1-3) times as long as deep, its length 
11-4-17-6 (M = 15-4) per cent of standard length. 



238 REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 

Mouth horizontal ; posterior maxillary tip reaching the vertical from the anterior 
orbital margin or extending somewhat beyond. Jaws equal anteriorly, the lower 
short and broad, from 30-0-37-8 (M = 35-6) per cent of head length, its length/ 
breadth ratio 1-0-1-4 (mode 1-2). H. simotes holotype is unusual in having its lower 
jaw only 28 per cent of the head length. 

Gill rakers short, 8 or 9 (rarely 10) on the lower part of the first arch. 

Scales ctenoid ; lateral line interrupted, with 30 (f.i), 31 (f.12) 32 (f.28), or 
33 (f-3) scales. Cheek with 2 or 3 (rarely 4) series of scales ; 6 or 7 (less frequently 
5 or 5 1) between dorsal fin origin and the lateral line ; 7 or 8 (rarely 6 or 9) between 
pectoral and pelvic fin insertions. 

Fins. Dorsal with 24 (f.7), 25 (f.33) or 26 (f.4) rays, anal n (f.7), 12 (f.32) 
or 13 (f.5), comprising XIV-XVIII, 8-10 and III, 8-10 spinous and soft rays. First 
pelvic ray produced, extending to the vent or even to the soft anal ; its posterior 
extension not correlated with sex or maturity. Pectoral fin shorter than the head. 
Caudal sub-truncate or feebly rounded ; scaled on the proximal half to two-thirds. 

Teeth. The outer series is composed of close set, movably implanted bicuspid 
teeth, with long, slender necks and expanded crowns. Cusp size in some individuals 
is markedly disparate, whilst in others the cusps are sub-equal. In the upper jaw, 
teeth situated postero-laterally are either tri- or unicuspid. 

A weak positive correlation exists between the number of teeth in the outer series 
of the upper jaw and standard length. 



S.L. (mm.) . 


• 49-58 


. 59-68 , 


, 69-78 


. 79-93 


Tooth number 


. 40-50 


46-60 


46-60 


■ 54-70 


Mean . 


. 46 


- 52 


52 


62 


N 


19 


6 


. 14 


4 



The inner series is composed of small tricuspid teeth ; 3-7 (mode 4) rows in each 
jaw. Compared with other Lake Victoria Haplochromis (except some individuals 
of H. nuchisquamulatus and H. lividus) the space separating inner and outer tooth 
series is greatly reduced in H. nigricans ; it is non-existent in 30 per cent of the 
specimens examined. 

Lower pharyngeal bone sub-equilaterally triangular ; dentigerous surface some- 
what broader than long ; teeth numerous, and similar to those in H. obliquidens 
and H. lividus. 

Cranial skeleton. The short and strongly decurved snout is reflected in the 
neurocranial shape. This differs slightly from that of generalized Haplochromis by 
having a more steeply sloping ethmo-vomer complex. Also, the dentary is relatively 
stouter and more massive in H. nigricans. 

Coloration in life : Breeding males. Ground colour black, shot with metallic 
blue ; snout, lips, interorbital region and to a lesser degree, cheeks and opercula, 
bluish. Dorsal fin black, lappets and maculae on the soft part deep crimson ; anal 
dusky crimson, ocelli yellow ; caudal crimson, pel vies black. Adult females and 
juveniles. Ground colour olivaceous ; a faint golden-yellow flush over the opercula 
and branchiostegal membrane. Dorsal and anal fins dark yellow ; caudal grey- 
green ; pelvics dusky yellow. 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 239 

Preserved material : Adult males. Black or slate grey ; seven or eight trans- 
verse bars visible on the flanks of light coloured fishes. Dorsal fin black, with pale 
margin and maculae ; caudal black proximally, pale distally ; anal pale ; pelvics 
black. Females and juveniles. Ground colour greyish-brown, with seven or eight 
dark transverse bars on the flank ; a pronounced lachrymal stripe. All fins hyaline 
or slightly darkened. 

Particular interest attaches to a single adult female with black and yellow piebald 
coloration similar to that described in Macropleurodus bicolor (Blgr.) and Hoplotilapia 
retrodens Hilgen. (Greenwood, 1956). The significance of this atypical individual is 
difficult to assess. In other characters H. nigricans does not manifest any apparent 
relationship with the monotypic genera, nor with H. sauvagei (Pfeffer), another 
species exhibiting sex-limited polychromism. It is probably the result of inde- 
pendent but parallel mutation occurring in H. nigricans, and therefore of no phyletic 
value. Such a phenomenon might be expected amongst members of a recently 
evolved and oligophyletic species flock. 

Distribution. Lake Victoria and the Victoria Nile. Although most localities 
represented in the present collection are in Uganda, this should not be taken to 
indicate that H. nigricans is confined to, or more abundant in, these waters. The 
species has been seen in many areas, but its lithophilic habits render capture difficult 
except by unconventional or specialized gear. 

Numerous specimens have been caught at Godziba Island (i° 20/ S., 32 36' E.). 
This small, rocky outcrop lies slightly south and west of the centre of Lake Victoria 
and is distant from either the mainland or other off-shore islands. Because there is 
no indication of H. nigricans ever occurring in deep or sub-littoral waters, one is led 
to suppose that Godziba fishes are at present isolated from coastal populations, and 
have been isolated for some considerable time. With this in mind, the Godziba 
sample was carefully compared with others from the mainland, but no phenotypic 
peculiarities could be detected. 

Ecology : Habitat. H. nigricans is apparently confined to rocky and shallow 
areas of the littoral zone. Since rock exposures are not infrequent in the exposed 
littoral, its habitat, broadly speaking, overlaps that occupied by other algal-grazing 
Haplochromis species. No data are available for populations living in the Nile. 

Food. The intestine is long (ca. 2J-3 times S.L.) and coiled. Observations on 
fishes in the lake indicate that H. nigricans feeds by grazing on algae from rock 
surfaces, a conclusion which is supported by stomach content analyses. 

Ingested material from thirty-two stomachs showed a preponderance of diatoms 
over all other material. Specific identification of these plants was impossible, but 
the genera represented (chiefly Navicula, Synedra, Rhopalodia and Gomphonema) 
are typically epilithic or epiphytic in Lake Victoria (Ross, 1954). The absence, 
except from two stomachs, of fragmentary phanerogam tissue (an important element 
in stomach contents of other algal grazing species) was noteworthy, but explicable 
if H. nigricans graze from rock surfaces. 

Filamentous green algae (Spirogyra and Oedogonium) and blue-green algae occurred 
less frequently, and were apparently undigested. 

Very fine, sand-grain-like particles were recorded from thirteen stomachs. That 



240 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 



these might have been fragments derived from rock surfaces and not the bottom 
seems likely in the absence of bottom debris typically associated with a sand 
substrate. 

Breeding. Spawning sites are unknown. Courtship activity has been observed 
amongst fishes living over rocks near the Ripon Falls, but actual spawning was not 
seen. Two females have been found with embryos and larvae in the buccal cavity ; 
it is assumed that H. nigricans, like the generality of Haplochromis species, is a 
mouth-brooder. 

The smallest adult fishes recorded were a female 51 mm. and a male 55mm. in 
standard length. Males apparently reach a larger size than females since no 
female greater than 70 mm. S.L. has been captured. 

Affinities. Haplochromis nigricans is closely related to H. serridens Regan of 
Lake Edward [vide Trewavas, 1933). Both species have almost identical dental 
morphology and pattern, as well as similarity in general facies and preserved colora- 
tion. No clear-cut quantitative characters can be found to separate the species. 
There is, however, a subtle difference in their gross morphology, probably attribu- 
table to the more rounded physiognomy of H. serridens. Also, the inner tooth bands 
of this species are usually broader and possess more teeth than those of H. nigricans. 

Tooth form, and less obviously the dental pattern, in one other Lake Edward 
species, H. fuscus Regan, is similar to that of H. nigricans ; but the species are 
readily distinguished by the smaller nuchal and thoracic scales in H. fuscus and also 
by its thicker lips and more abruptly declivous dorsal head profile. 

Amongst Lake Victoria species H. nigricans is probably related to, and derived 
from a species resembling H. nuchisquamulatus. 

Diagnosis. H. nigricans is distinguished from other Lake Victoria Haplochromis 
with bicuspid outer teeth by the following combination of characters : a short and 
broad lower jaw (modal length /breadth ratio 1:2); slender, movably implanted 
outer teeth narrowly separated, if at all, from the broad bands of inner teeth ; a 
strongly decurved dorsal head profile ; a long and convoluted intestine. 



Study material and distribution records 

Museum and reg. no. 

British Museum (N.H.) 1906.5.30.469 (Holotype 
of Tilapia nigricans) .... 

Genoa Museum (Holotype of Tilapia simotes) . 

British Museum (N.H.) 191 1.3.3. 160-163 (Para- 
types of T. simotes) ..... 

British Museum (N.H.) 191 1 . 3.3. 156-158, plus 
one additional specimen (Paratypes of T. simotes) 

British Museum (N.H.) 1956.7.9. 129-136 

„ „ „ 137-148 
„ „ „ 149-15° 



151 



152 
153-165 



Locality. 

Entebbe 

Kakindu (Victoria Nile) 

Jinja (Ripon Falls) 

Kakindu 

Napoleon Gulf, near 

Ripon Falls 
Jinja Pier 
Napoleon Gulf, near 

Jinja 
Beach near Nasu Point, 

Buvuma Channel 
Buka Bay (Uganda) 
Godziba Island 



Collector. 

Degen. 
Bayon. 

Bayon 

Bayon 
E.A.F.R.O. 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 241 

Haplochromis nuchisquamulatus (Hilgendorf) 1888 

Chromis nuchisquamulatus Hilgend., 1888, 5. B. Ges. naturf. Fr. Berlin, 76. 

Ctenochromis nuchisquamulatus (Hilgend.), Pfeffer, 1896, Thierw. O. Afr. Fische, 14. 

Tilapia nigricans (part), Blgr., 1915, Cat. Afr. Fish., 3, 241. 

Haplochromis nuchisquamulatus (part), idem, ibid., 290. 

Haplochromis (Neochromis) nuchisquamulatus (Hilgend.), Regan, 1922, Proc. zool. Soc, London, 

163. 

The holotype of H. nuchisquamulatus is amongst those specimens, once housed in 
the Berlin Museum, which cannot be located at the present time. It is thus the more 
regrettable that Hilgendorf 's original description is totally inadequate for modern 
taxonomic purposes. 

As a basis for comparison I have therefore relied upon Regan's identification of 
two British Museum (Nat. Hist.) specimens. From Regan's paper (1922) it is clear 
that he, too, was unable to study the type specimen, but he apparently gained 
sufficient information from photographs and data supplied by Dr. Pappenheim to 
identify his material. Of this I have located only one specimen (British Museum 
(N.H.) reg. no. 1911.3.3.155, from Kakindu, Victoria Nile). In its general 
morphology this fish agrees closely with a photograph of the type. Further, with 
the aid of a binocular microscope it has proved possible to check certain other 
characters visible in this remarkably clear photograph. 1 

Although this species is represented in my study-material by only six specimens, 
I have little doubt as to its biological validity. Morphologically H. nuchisquamulatus 
is intermediate between H. lividus and H. nigricans : it may well represent the 
stock from which these species diverged. The tooth form of H. nuchisquamulatus 
is less specialized than that of //. lividus and is nearer H. nigricans. That is to say, 
the outer teeth are slender, bicuspid and movable, whilst those of the inner series 
show a tendency towards an increase in the number of rows and a decrease in the 
space separating them from the outer series. The lower jaw is more slender than in 
H. nigricans and is similar to the dentary in H. lividus and H. obliquidens, and in 
other, more generalized Haplochromis. 

None of the dental and associated characters considered above lies within the 
known range of intra-specific variability for H. nigricans or H. lividus. Neither is 
there any indication by analogy with well-defined Haplochromis species that the 
" nuchisquamulatus " character-complex is an extreme variant of some other species 

Description. The principal morphometric characters for each of the six speci- 
mens examined are tabulated below. All are adult males. 

S.L. Depth.* Head.* Po. % Io. % Snt. % Eye. % Ch. % Lj. % C.P.* 



83-0 


38-0 


31*3 


15*4 


30-8 


30-8 


30-8 


23-0 


36-5 


15*7 


86-o 


37*2 


32-5 


14*3 


28-6 


28-6 


32-2 


23-2 


35'7 


15-2 


93 -o 


38-7 


32*3 


i4'5 


29*0 


32*2 


29*0 


24*2 


38-6 


i5*i 


98-0 


38-8 


3i-6 


16*2 


30-0 


32-2 


25-8 


22*6 


38-7 


15*3 


99. 


37'4 


32*8 


15*4 


30-8 


30-8 


30-8 


24*6 


40*0 


15-2 


113-0 


37*o 


3o-3 


17-4 


26*5 


3i-8 


29-0 


26-5 


36-2 


18-6 



* Percentage standard length. 
% Percentage head-length. 



1 To be reproduced in a later part of this series. 



242 REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 

Dorsal head profile curved and sloping. Mouth horizontal ; posterior maxillary- 
tip extending to the vertical from the anterior orbital margin or slightly beyond. 
Jaws equal anteriorly, the length /breadth ratio of the lower 1-4-17 (mode 1-5). 

Teeth. Outer teeth unequally bicuspid ; a few slender and unicuspid teeth 
occur posteriorly in the upper jaw, in which there are from 50-70 teeth. Although 
relatively fine, the neck in these teeth is stouter and less clearly demarcated from the 
expanded crown, than in H. lividus or H. obliquidens. 

Inner teeth small and tricuspid, occurring in 4-8 and 3-6 rows in the upper and 
lower jaws respectively ; the space separating inner and outer series is reduced. 

Lower pharyngeal bone sub-equilaterally triangular, its dentigerous surface 
slightly broader than long. Teeth fine and numerous ; in the three larger specimens, 
the median teeth are enlarged. 

Gill rakers short, 8-10 on the lower part of the first arch. 

Scales ctenoid : lateral line interrupted, with 31 (f.2), 32 (f.2) or 33 (f.2) scales. 
Cheek with 2 or 3 series. 6-8 scales between dorsal fin origin and the lateral line ; 
6-8 scales between pectoral and pelvic fin insertions. 

Fins. Dorsal with 24 (f.i), 25 (f.2) or 26 (f.3) rays, anal 12 (f.4) or 13 (f.2), 
comprising XV-XVII, 9 or 10 and III, 9 or 10 spinous and soft rays. First pelvic 
ray produced, extending to the second anal ray. Pectoral fins slightly shorter than 
the head. Caudal sub-truncate. 

Skeleton. That of a generalized Haplochromis. 

Coloration. Unknown in life and known only for preserved males. Ground 
colour dark greyish-brown, the dorsal and ventral surfaces darker than the 
flanks, across which seven transverse bars are visible ; in two specimens the chest is 
black. Well-defined, narrow lachrymal and two interorbital stripes ; two broad 
bands across the nape, one immediately post-ocular in position, the other slightly 
more posterior. 

Ecology. Of the six specimens studied, five were caught in exposed littoral 
zones of Lake Victoria, and one in the Victoria Nile. The type specimen is from 
Lake Victoria, but no precise locality is given. 

Food. Fragments of plant tissue and numerous epiphytic algae were recorded 
from four of the five stomachs examined, whilst the fifth contained filaments of 
Oedogonium and some fragmentary plant tissue. 

Diagnosis. H. nuchisquamulatus is distinguished from other Lake Victoria 
Haplochromis with bicuspid outer teeth by the following combination of characters : 
long and convoluted intestine (ca. 3 X S.L.) ; relatively slender, movably implanted 
and numerous outer teeth ; increased number of inner tooth rows (3-8) narrowly 
separated from the outer series. From H. nigricans it is recognized by the narrower 
lower jaw and less strongly decurved dorsal head profile ; outer teeth in H. nuchi- 
squamulatus are also somewhat stouter than those of H. nigricans. These acutely 
cuspidate teeth serve to separate H. nuchisquamulatus from H. lividus. 

The diagnostic character used by Hilgendorf (small nuchal scales whose exposed 
surface is less than half that of flank scales) cannot be considered valid. In most 
Haplochromis nuchal scales are smaller than those on the flank, and furthermore are 
subject to quite considerable intra-specific size- variation. 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 

Study material and distribution records 

Museum and reg. no. 

British Museum (N.H.) 1911.3.3.154 
British Museum (N.H.) 1906.5 .30.316-317 
British Museum (N.H.) 1956.7.9. 166-167 



243 



168 



Locality. 

Kakindu (Victoria Nile) 

Entebbe 

Beach near Nasu Point, 

Buvuma Channel 
Godziba Island 



Collector. 

Bayon. 
Degen. 

E.A.F.R.O. 
E.A.F.R.O. 



SUMMARY 

1. The algal-grazing species Haplochromis obliquidens Hilgendorf 1888, H. 
nigricans (Boulenger) 1906, and H. nuchisquamulatus (Hilgendorf) 1888, are re- 
described on the basis of new and more extensive collections. 

2. A new species, H. lividus, apparently related to H. obliquidens is described. 

3. Data on the food and ecology of these species are given. 

4. Consideration is given to the possibility of recognizing a number of supra- 
specific groups of Haplochromis in Lake Victoria. At present, although such groups 
may be determined, it is impossible to give them formal taxonomic status. 



ACKNOWLEDGMENTS 

I wish to acknowledge my gratitude and thanks to the Trustees of the British 
Museum (Natural History) for facilities afforded me during the tenure of a Colonial 
Fisheries Research Studentship ; to Professor L. Bertin of the Museum National 
d'Histoire naturelle, Paris, for allowing me to study type specimens of Lake Victoria 
Cichlidae described by Pellegrin ; to Dr. Delfa Guiglia of the Museo Civico di 
Storia Naturale, Genoa, for the courtesies mentioned in the text ; to my colleague 
Dr. Philip S. Corbet for identifying some of the material from gut contents ; and to 
Mr. Denys W. Tucker, for his very helpful criticism of the manuscript. I am 
especially indebted to Dr. Ethelwynn Trewavas for much helpful information 
and advice. 



REFERENCES 
(Other than those given in full in the text) 

Baerends, G. P. & Baerends Van Roon, J. M. 1950. An introduction to the study of the 
ethology of cichlid fishes. Behaviour, Supplement 1, pp. 1-242. Leiden. 

Greenwood, P. H. 1951. Evolution of the African cichlid fishes ; the Haplochromis species 
flock in Lake Victoria. Nature, London, 167 : 19. 

1956. The monotypic genera of cichlid fishes in Lake Victoria. Bull. Br. Mus. nat. 

Hist., Zool. 3, no. 7. 

Pfeffer, G. 1893. Ostafrikanische Fische gesammelt von Herrn Dr. F. Stuhlmann. Jahrb. 
Hamburg, wiss. Anst. 10 (2), 129-177, 3 pis. (1-49 sep. pag.). 

Poll, M. & Damas, H. 1939. Poissons. Exploration du Pare National Albert, mission H. 
Damas (1935-1936), fasc. 6, pp. 1-73. 

Regan, C. T. 1920. The classification of the fishes of the family Cichlidae. I. The Tangan- 
yika genera. Ann. Mag. nat. Hist. (9) 5 : 33. 



244 



REVISION OF LAKE VICTORIA HAPLOCHROMIS SPECIES 



Regan, C. T. 1921. The cichlid fishes of Lakes Albert Edward and Kivu. Ann. Mag. nat. Hist. 

(9) 8 : 632. 
— ■ — ■ 1922. The cichlid fishes of Lake Victoria. Proc. zool. Soc. Lond. 157. 
Ross, R. 1954. The algae of the East African Great Lakes. Proc. International Assoc. : 

Theoretical and Applied Limnology . 
Trewavas, E. 1933. Scientific results of the Cambridge expedition to the East African lakes, 

1 930-1. II. The cichlid fishes. /. Linn. Soc. {Zool.) 38 : 309. 
— ■ — ■ 1935. A synopsis of the cichlid fishes of Lake Nyasa. Ann. Mag. nat. Hist. (10) 16 : 65. 
Worthington, E. B. 1932. A Report on the Fisheries of Uganda. Crown Agents, London. 




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A SYSTEMATIC REVISION 

OF THE FISHES OF THE TELEOST 

FAMILY CARAPIDAE 

(PERCOMORPHI, BLENNIOIDEA), 

WITH DESCRIPTIONS OF TWO 

NEW SPECIES 



D. C. ARNOLD 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 4 No. 6 

LONDON: 1956 



A SYSTEMATIC REVISION OF THE FISHES 

OF THE TELEOST FAMILY CARAPIDAE 

(PERCOMORPHI, BLENNIOIDEA), 

WITH DESCRIPTIONS OF TWO NEW SPECIES 



BY 

D. C. ARNOLD 

(Gatty Marine Laboratory, and Department of Natural History, St. Andrews) 



Pp. 245-307 ; 20 Text-figures. 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ZOOLOGY Vol. 4 No. 6 

LONDON : 1956 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 4, No. 6 of the Zoological series. 




PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued November, 1956 Price One Pound 



A SYSTEMATIC REVISION OF THE FISHES 

OF THE TELEOST FAMILY CARAPIDAE 

(PERCOMORPHI, BLENNIOIDEA), 

WITH DESCRIPTIONS OF TWO NEW SPECIES 

By D. C. ARNOLD 

SYNOPSIS 

The life history, mode of life and behaviour of Carapus acus (Briinnich) are briefly reviewed 
and compared with those of other species. An account of the range of skeletal structure within 
the Carapidae is given as a basis for generic separation, the various species are redescribed and 
their synonymies are revised. Two new species of the genus Carapus are described. Keys are 
given for the diagnosis of the genera, subgenera and species recognized. The paper is illustrated 
by 20 text-figures and includes a list of the principal references to the family Carapidae. 

CONTENTS 

Page 

I. Introduction . . . . . . . . . 247 

II. The Life History of Carapus acus ...... 248 

III. Behaviour and Mode of Life ....... 252 

IV. Structure ........... 254 

V. Systematic Arrangement ........ 259 

1. Genus Carapus ......... 260 

2. Genus Echiodon ......... 288 

3. Genus Encheliophis ........ 295 

VI. References Cited ......... 302 

I. INTRODUCTION 

The best known member of the Carapidae is the Mediterranean species Carapus 
acus (Briinnich), a monograph on which was published over seventy years ago 
(Emery, 1880). This account was principally devoted to the adult anatomy, but 
also included observations on the life history and behaviour of the fish. Other 
members of the family have been briefly recorded in a number of ichthyological 
works, but no general account of the family appears ever to have been published. 
The work here recorded was performed mainly in Italy and at Plymouth. Studies 
on living fish were undertaken during tenure of the Oxford and Royal Society Tables 
at the Stazione Zoologica, Naples, while the taxonomic portion of the work was 
performed almost entirely during tenure of the Oxford Table at the Laboratory of 
the Marine Biological Association of the United Kingdom, Plymouth. Short periods 
were also spent in study at the British Museum (Natural History) and at the Scottish 
Home Office Laboratory, Aberdeen. 

zool. 4. 6. 17 



248 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

Preserved specimens were obtained from certain other sources, notably from the 
following institutes : 

Universitetets Zoologiske Museum, Copenhagen. 
Institut Oceanographique, Monaco. 
Museu Municipal, Funchal, Madeira. 
Institut za Oceanografiju i Ribarstvo, Split. 

Information on the distribution of various species was obtained by correspondence 
with the following laboratories : 

Institut d'Hydrobiologie et de Peche, Alexandria, Egypt. 

Marine Biological Station, al Gardaqa, Egypt. 

Institut Scientifique Cherifien, Rabat, Morocco. 

Laboratorio Oceanografico, Malaga, Spain. 

Station Biologique, Roscoff, France, 

Station Biologique, Archachon, France. 

Institut za Biologiju Mora, Rovinj, Yugoslavia. 

Zoology Department, University College, Achimota, Gold Coast. 

East African Marine Fisheries Research Organization, Zanzibar. 

The University, Travancore, India. 

Fisheries Department, Penang, Malaya. 

Department of Agriculture, Sandakan, North Borneo. 

Department of Harbours and Marine, Brisbane, Australia. 

Department of Zoology, The University, Sydney, Australia. 

Victoria University College, Wellington, New Zealand. 

Department of Zoology, The Museum, Dunedin, New Zealand. 

Radiographs of representative specimens were made by the Radiography Depart- 
ment of the Royal Naval Hospital, Devonport, and of type material by Mr. A. C. 
Wheeler of the British Museum (Natural History) . 

My most grateful thanks are due to the Directors and Staffs of these various 
institutions, without whose kindly co-operation and assistance this work would not 
have been possible and to the many individual workers with whom I have discussed 
the results of this investigation and the problems arising therefrom. 

During the course of this study financial support was provided by the Oxford 
Naples Scholarship and by a maintenance grant and supplementary grants from the 
Department of Scientific and Industrial Research. 

II. THE LIFE HISTORY OF CARAPUS ACUS 

Eggs attributable to C. acus are found in the Mediterranean during July, August, 
and early September, floating at the sea surface in yellowish, oval masses each 
containing some thousands of eggs. The egg is ellipsoidal, with diameters of 0-90 
mm. and 0*75 mm., and has a large oil-globule, the yellowish tint of which is 
responsible for the colour of the egg-rafts (Raffaele, 1888). Spawning has not been 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 249 

observed, not has artificial fertilization been achieved. Embryonic development is 
rapid, the larva hatching in an anatomically incomplete condition on the third day 
after spawning. Emery described the early development in general terms, but owing 
to the paucity of material the embryology of the fierasfers has never been subjected 
to critical study by modern methods. 

At hatching, the young fish enters the first larval stage, the vexillifer, characterized 
by a long dorsal appendage, the vexillum. On the first day of post-embryonic 
life this appendage is a small, pigmented thickening at the anterior end of the dorsal 
fin, but as the larva grows the thickening enlarges, forming first a small papilla, 
then a soft, forked projection and finally a long, lobed structure whose size and 
complexity increase with the increasing size of the larva (Emery, 1880). As the 
vexillum grows, its pigment is limited first to the lobes and then to their proximal 
portions, leaving the stalk and distal parts of the lobes relatively unpigmented in the 
older larvae. Later still the vexillum degenerates and the larva sinks from the 
surface into the deeper layers until finally, with the appendage reduced to a mere 
projection or even lost entirely, the young fish enters upon the benthic mode of life 
of the second carapid larva. It is probable that regression does not often run its 
full course, for the fragility of the vexillum is such that few fish are obtained with it 
undamaged and it must often be broken off instead of resorbed. 

Emery hatched fierasfer larvae in the aquarium and reared them for about a week, 
but was unable to feed the fish and could not follow their development after they had 
attained a length of 3-4 mm. Further knowledge of the growth and metamorphosis 
of the vexillifer has been obtained solely from specimens taken in plankton hauls, 
most of which were never studied alive. The youngest vexillifers taken in the 
plankton were a pair obtained by Emery. They were about 10 mm. long, but showed 
little advance on those he was able to rear. Others studied by Padoa (1947) ranged 
from 15 mm. to 85 mm. in length. From this author's account it appears that 
increase in size is unaccompanied by any great changes in proportion. The head 
remains approximately one-fifteenth of the total length, the preanal length about 
one-tenth of the total, while throughout the vexillifer stage the end of the abdominal 
cavity is only a short distance behind the anus. A 76 mm. vexillifer mentioned by 
Emery agrees well with Padoa' s series, as do the large specimens described by 
Gasco (1870) and Costa (1871) under the name Vexillifer dephilippii Gasco. 

The second larval stage of C. acus is at first benthic, later inquiline. It was 
originally described as Encheliophis tenuis Putnam (1874) and Padoa (1947) has 
proposed that it be termed the tenuis larva. The tenuis has been found far less 
frequently than has the vexillifer and records of perhaps not more than 20 have 
appeared in the literature. 

The tenuis stage is essentially a phase of growth and change. It lacks the vexillum 
and is characterized principally by the immense length of the cylindrical body and 
the relative smallness of the head. In the later vexillifers the head comprises about 
one-fifteenth of the total length, yet in the youngest tenuis so far described (Emery, 
1880) the head constituted but one-thirtieth of the total and its actual length, 
5 mm., was no greater than that of the head of a vexillifer of only half its size. 

While the tail is elongating, the head and trunk of the tenuis are also growing, 



250 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

though not nearly so rapidly. Thus the disparity between head and trunk length 
on the one hand and total length on the other continually increases. At a total 
length of about 200 mm. further changes occur. Elongation is now replaced by 
shortening ; tail growth by tail resorption. But though the tail region of the tenuis 
now decreases in length, the head and trunk regions continue to grow and the propor- 
tions of the body approach ever more closely to those of the adult fish. However, 
even in the anterior region of the body not all parts develop alike, for while the head 
and abdominal cavity elongate, the anus remains at about the same distance from 
the snout and thus occupies an ever more advanced position. During this and 
subsequent stages of development the depth of the body also increases, though the 
width remains fairly constant, and the cylindrical body of the tenuis is converted to 
the compressed form of the adult fierasfer. 

The tenuis lacks pigment except for the silvery iris and black and red chromato- 
phores on the top of the head. Trunk and tail are glassily transparent. When 
removed from a holothurian, the tenuis has a pinkish tinge which fades after a short 
time in sea water and is never recovered. It is apparently due to the colour of the 
blood. 

The tenuis does not metamorphose directly into the adult, but into a well-marked 
juvenile stage (Arnold, 1953). This is primarily a period of consolidation in which 
the developmental trends of the tenuis are continued to produce the adult form. 
At the close of metamorphosis the head of the young fish is about one-twelfth of the 
total length ; the anus is beneath, or only a little behind, the vertical through the 
roots of the pectoral fins ; the pectorals are round and short, one-quarter to one-third 
the length of the head ; and patches of black pigment have begun to appear on the 
still translucent tail. 

At first the total length, 70-80 mm., alters but little, though the head continues 
to elongate and the trunk to deepen. When the adult proportions have been nearly 
obtained and the length of the head is between one-seventh and one-eighth of the 
total length, the fish elongates once more. Now, however, all parts of the body 
grow at about the same rate and there is little further alteration in proportions, 
beyond a deepening of the anterior part of the trunk and the advancement of the 
anus to a position a little in front of the pectoral fins. The changes in length of 
head relative to total length during the life history of C. acus are shown graphically 
in Text-fig. 1. As growth proceeds, the pectorals elongate, becoming oval and about 
one-half the length of the head, while red chromatophores appear among the black. 
The reddish tinge given to the body marks the end of the juvenile stage and may 
perhaps be associated with the onset of sexual maturity. Behavioural changes also 
occur at this time. 

The dentition of the developing C. acus, owing to the importance of dental 
characters in the identification of adult fierasfers, deserves special consideration. 
The smallest vexillifers lack teeth, but in later individuals teeth appear on jaws, 
palatines and vomer. These teeth are at first uniserial in arrangement and of 
uniform size, but as the vexillifer grows a pair of rather larger teeth appears at the 
front of the upper jaw, followed by a similar pair in the lower jaw, and both pairs are 
fully developed by the time that a total length of 60 mm. has been attained (Padoa, 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 251 



200- 




100- 



Length of Head 



Fig. 1. — Changes in proportions (head length relative to total length) during the life 
history of Carapus acus. a, Vexillifer ; b, early tenuis ; c, late tenuis ; d, juvenile ; 
and e, adult stages. 



252 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

1947). Towards the end of the vexillifer stage the size difference between these 
anterior teeth and those in the rest of the jaw diminishes and towards the close of 
the tenuis stage enlarged anterior teeth are no longer present. Throughout the 
tenuis period the dentition remains uniserial on jaws and palatines, but at the 
beginning of the juvenile stage the number of rows begins to increase until the com- 
pletely polyserial condition of the adult is attained. 

It has been suggested (Dean, 1895) that the elongated bodies of the fierasfers, 
their persistent notochord and functional pronephros are expressions of a general 
paedomorphic tendency of the family and it is probable that the dental characters 
of many species, the enlarged anterior jaw teeth of certain Carapus spp. and the 
uniserial dentition of Encheliophis, are also larval characters perpetuated in the adult 
form. In the circumstances, considerable doubt is cast upon the validity of the 
dentition as a diagnostic character for the separation of new species unless it can 
clearly be shown that the types are mature individuals, while the occurrence of 
partially uniserial dentition in a fierasfer of small size can probably always be taken 
as indicating that the specimen in question is in the juvenile stage of its development. 



III. BEHAVIOUR AND MODE OF LIFE 

The behaviour of the adults of Carapus acus was first studied by Emery (1880), 
who found that the fish swam in a sharply-tilted, head-down position and entered 
the host holothurian tail-first through the anus. Recent observations on the same 
species (Arnold, 1953) have shown pronounced behavioural differences between fish 
in different stages of the life history. The youngest specimens of C. acus studied 
alive were tenuis larvae obtained from Holothuria tubulosa. When removed from 
the host and placed in sea water they showed a characteristically violent movement. 
The body was slanted upwards, the head even thrust above the surface of the water, 
and its flexures were of extremely wide amplitude, very different in appearance from 
the normal swimming movements. Such " tenuis " movements were occasionally 
shown by juveniles and adults when removed from their hosts or when attempting 
to evade capture. Fish displaying " tenuis " movements did not respond in any 
way to the presence of a holothurian, nor did the tenuis larvae themselves appear 
to be capable of entering a new host. Juveniles and adults, however, responded at 
once to the presence of a holothurian and would swim the length of its body until 
they located the anus, then attempt to enter. Adults almost always entered tail- 
first by means of a pronounced corkscrew motion in which the body of the fish rotated 
through 360 or more. The juveniles normally entered head-first, though some of 
the older specimens might make incomplete and unsuccessful attempts at a tail- 
first entry. Only those fish which had attained their full growth in the juvenile 
condition and were beginning to assume the form and colour of the adult displayed 
both modes of entry. 

Behaviour differences such as these may be shown also by other fierasfers, but at 
present the few recorded observations on other members of the Carapidae relate 
almost entirely to adult fish. Tail-first entry has been described for the adults of 
C. bermudensis (Jones) (Linton, 1907 ; Aronson & Mosher, 195 1) and C. hornet 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 253 

(Richardson) (Mukerji, 1937). The entry of Encheliophis gracilis (Bleeker) into 
starfish has been observed (Doleschall, 1861), but it is not clear from this account 
whether entry was head-first or tail-first. Encheliophis hancocki (Reid) has been 
observed both to enter and leave its host head-first (Steinbeck & Ricketts, 1941). 
Nothing whatsoever is known of the behaviour of those species which inhabit 
lamellibranchs. 

Most accounts of the behaviour of fierasfers have been purely descriptive and little 
attempt has been made to analyse the behaviour of any species in terms of stimuli 
and responses. Studies on adults and juveniles of C. acus by means of various 
models showed that the fish responded only if the water contained mucus from a 
holothurian and provided that the model was long relative to its depth (Arnold, 
unpublished observations). In the absence of a chemical stimulus C. acus shows 
none of the exploratory movements that usually precede attempt at entry, while 
an ovoid or circular model is usually ignored. Presence of a water current seems to 
be a necessary prerequisite for actual penetration, though in the absence of prior 
chemical and visual stimuli a water current alone has either no effect or is actually 
repellent to the fish. Recent work on C. bermudensis has shown that in this species 
chemical and tactile stimuli are of most importance (Aronson & Mosher, 1951), 
but a full account of this work has yet to be published. 

C. acus lies within the body cavity of its host, usually at the anterior end among 
the branches of the gonads, on which it apparently feeds. The tenuis larva does 
not survive long after removal from its host and the fierasfer must thus remain 
within its first holothurian throughout this period of its life history. Juveniles and 
adults live well in sea water and small Crustacea have been found among their 
stomach contents. It is probable that C. acus leaves its host only when the 
holothurian eviscerates. 

Little is known of the location of other fierasfers within the bodies of their hosts. 
Encheliophis gracilis apparently breaks into the body cavity of the starfish it in- 
inhabits (Doleschall, 1861 ; Yosii, 1928), and Encheliophis vermicularis Muller has 
been found within the body cavity of its host holothurian, feeding upon the viscera 
(Semper, 1861). Both Encheliophis hancocki and C. bermudensis have been observed 
freely to enter and leave their hosts (Steinbeck & Ricketts, 1941 ; Linton, 1907) 
and thus can hardly have penetrated further than the cloaca or base of the branchial 
trees. Different species of fierasfer doubtless differ in their dependence upon their 
hosts and in the distance to which they penetrate into the body. 

Some fierasfers seem to show little host specificity. C. homei, for example, has 
been recorded from holothurians, asteroids, echinoids, lamellibranchs and a tunicate. 
Others however, are restricted to particular host species. Thus C. bermudensis has 
been recorded only in Actinopyga agassizi, Encheliophis sagamianus (Tanaka) only 
in Holothuria monacaria, and C. acus only in H. tubulosa and Stichopus regalis. In 
the laboratory C. acus also enters H. poli, H. helleri and H, sanctori, but does not 
respond to Cucumaria planci or Phylloporus urna. It does not penetrate into the 
body cavity of H. poli, but lodges in the branchial trees and usually emerges within 
24 hours. Choice experiments did not provide evidence that C. acus could 
distinguish between H. tubulosa and H. poli prior to entry. 



254 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 



IV. STRUCTURE 

A full account of the structure of the commonest European fierasfer, Carapus 
acus, was given by Emery (1880), but little is known of the anatomy of any other 
species. Owing to the general rarity of the fierasfers it has been impossible to make 
a comparative study of their soft anatomy, but X-ray examination of a number of 
the types and of other clearly identifiable specimens has enabled the skeletons of 
a number of species to be studied without damage to the specimens themselves. 
In all species examined the tail and posterior trunk vertebrae have been found to 
possess the same general characters, differing only in size and relative degree of 
ossification. The centra are characteristically hour-glass shaped, generally twice as 
long as wide or high, and surmounted by long, pointed, backwardly directed neural 
spines which in the trunk region project above the succeeding vertebrae, but in the 
tail region become progressively smaller until towards the tip of the tail they are no 
longer recognizable. The transverse processes of the trunk vertebrae jut out at 
right angles from the centra, then turn downward and backward. The haemal 
spines of the tail vertebrae are extremely long anteriorly, but decrease in height 
towards the tip of the tail. In this region ossification becomes so slight that it is 
usually impossible to count the total number of vertebrae, the most posterior of 
which are mere rings encircling the persistent notochord. 

But though the various species resemble each other in the form of the majority of 
their vertebrae, in the structure of the anterior vertebrae and of the lower jaw they 
are classifiable into three sharply separated groups, two of which may be further 
subdivided by means of other characters. In the systematic portion of this account, 
this subdivision of the Carapidae on the basis of skeletal characters has been used as 
the basis for generic separation of the adults. Though it has not been possible to 
examine the skeletal characters of larval fierasfers, the lower jaws (which can easily 
be seen by superficial examination) show similiar variations to the lower jaws of 
adults and thus provide grounds for generic classification even of immature specimens. 

The majority of fierasfers examined — including representatives of the species 
originally described as Gymnotus acus Brunnich, Fierasfer dubius Putnam, Lefroyia 
bermudensis Jones, Oxybeles Homei Richardson, Fierasfer parvipinnis Kaup, Fie- 
rasfer affinis Gunther, Fierasfer caninus Gunther, Fierasfer mar gar iti ferae Rendahl 
and Carapus parvibrachium Fowler — form a fairly homogenous group which in the 
main corresponds with the genus Carapus as at present understood. The majority 
of these species have 17-18 trunk vertebrae (19-20 in Fierasfer mar gariti ferae and 
Carapus parvibrachium), the first of which has a roughly cubical centrum surmounted 
by a stout neural spine with a truncated tip. This spine slopes forward and appears 
to be attached to the skull by a calcified ligament. The transverse processes of the 
first vertebra are slender and scimitar-shaped, directed backward and downward. 
The second vertebra resembles the first in the form of its centrum and transverse 
processes, but has a pointed neural spine which slopes slightly backward. The 
third and fourth vertebrae have hour-glass shaped centra, long backwardly-projecting 
neural spines and expanded transverse processes which are fused together into broad 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 255 




Fig. 2. — Vertebrae of Carapus acus. a, Nos. i and 2, lateral aspect (transverse processes 
omitted) ; b, nos. 1 and 2, dorsal aspect ; c, nos. 3 and 4, dorsal aspect ; d, nos. 3 and 4, 
lateral aspect ; e, representative posterior trunk vertebra ; f, representative anterior 
tail vertebra. {Camera lucida drawings from radiographs.) 



flat plates, rounded anteriorly and tapering posteriorly to points level with the middle 
of the fifth vertebra (Text-fig. 2). 

In this group of fierasfers the lower jaw is nearly flat along its lower edge. The 
upper tooth-bearing edge commences parallel to the lower, then curves upwards to 
the greatly expanded proximal portion of the jaw (Text-fig. 3 a) . The narrowest part 
of the jaw is the extreme tip, from which a lateral ridge, parallel to the lower edge, 
extends to the point of articulation with the cranium. Just beneath this ridge there 
is at the distal extremity of the jaw a deep and narrow notch. The teeth are in a 
narrow band (a single row only in the larvae) along the entire exposed surface of 
the jaw. Each tooth is separated from its neighbours by a space almost as wide as 
the diameter of the base of the teeth. 



256 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

Two of the species included within this group, Fierasfer mar gar iti ferae and 
Carapus parvibrachium, could not be examined in such detail as the rest owing to the 
presence of a large reniform, calcareous body lying in the midline in the anterior 
part of the body. This structure, presumably the anterior part of the swim-bladder, 
effectively concealed parts of the second, third and fourth vertebrae. So far as 
they could be seen, however, they appeared similar in general structure to those of 
of the other species, though undoubtedly differing in detail. These two species differ 






5 mm 

Fig. 3. — Jaw structure in the Carapidae. Lower jaws of a, Carapus ; b, Echiodon ; and 
and c, Encheliophis . (Semidiagrammatic drawings based upon radiographs.) 



also from the others in their slightly greater number of trunk vertebrae and in 
certain aspects of their dentition. In the systematic account they are therefore 
accorded subgeneric status. 

The second group of fierasf ers contains only two species which have been available 
for examination, those originally described as Ophidium dentatum Cuvier and 
Echiodon Drummondi Thompson. These species have been frequently confused with 
each other and generally assigned to Carapus. Both in structure and appearance, 
however, they differ considerably from the group of species so far considered. The 
body is considerably longer and the trunk region of the vertebral column comprises 
27-28 vertebrae, the first two of which are similar to the first two of the species 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 257 

already considered. The third and fourth vertebrae, in contrast, are entirely 
separate from each other and show not the least sign of fusion of the transverse 
processes (Text-fig. 4). The edges of the lower jaw are flat and parallel, the proximal 
portion is small and at its tip the jaw first narrows, then expands again as a small knob 
(Text-fig. 3B) . A lateral ridge runs from this knob, across the narrowed portion of 
the jaw and as far as the articulation with the skull. The dentition comprises two 
series. Anteriorly one or two immense, fang-like teeth occupy the terminal knob, 
while the neck is toothless and its position easily visible in the intact animal as a 
pronounced diastema. The rest of the jaw is occupied by a narrow band of teeth 
(or single row only in the larva) closely pressed together. 

Composing the third group of fierasfers is a number of species which, judged 
externally, appear to fall into two classes. Examples of the species originally 




3 mm 



Fig. 4. — Vertebrae of Echiodon drummondi. a, No. i, lateral aspect ; b, no. 1, dorsal 
aspect ; c, no. 2, lateral aspect ; and d, no. 2, dorsal aspect. (Camera lucid a drawings 
from radiographs.) 



described as Oxybeles gracilis Bleeker and Encheliophis vermicularis Muller have been 
studied. Of these, E. vermicularis lacks pectoral fins though the girdle is still 
present, while specimens of 0. gracilis are individually more robust and possess small 
pectorals. In vertebral and jaw structure, however, 0. gracilis and E. vermicularis 
are almost identical. There are 30-31 trunk vertebrae, the first two of which 
resemble those already described for the first group of fierasfers, with the exception 
that the neural spine of the second vertebra is rather shorter and stouter and is 
attached to the back of the skull by a structure which is apparently a slender, 
calcified ligament. The insertion of this ligament spreads on to the anterior part 
of the neural arch of the third vertebra (Text-fig. 5). It is characteristic of this 
third group of fierasfers that the third, fourth and fifth trunk vertebrae have their 
transverse processes expanded and fused into wide lateral wings, the outer edges of 
which are smoothly curved, while their points reach as far back as the level of 
articulation between the sixth and seventh vertebrae. 



258 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

0. gracilis and E. vermicularis differ from all other fierasfers in the slender, curved 
form of the lower jaw. The narrowest portion is at the tip, the proximal part is 
little expanded and the lateral ridge is little developed (Text-fig. 3c). Even in the 
adult the teeth are in a single row only and are widely separated from each other, the 




2 mm 

Fig. 5. — Vertebrae of Encheliophis (Jordanicus) gracilis, a, Nos. 1-5, lateral aspect ; b, nos. 
3-5, dorsal aspect. (Camera lucida drawings from radiographs.) 



distance between successive teeth being as much as two or three times the diameter 
of the tooth-base. 

Various characters have been used for the discrimination of species within the 
Carapidae, but those which have proved most generally useful for diagnosis are the 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 259 

dentition and certain proportions of the body. Owing to their soft bodies, fierasfers 
are liable to shrinkage and distortion on preservation and in the following systematic 
description all measurements have been made on specimens preserved either in 
formalin or, more usually, in alcohol. In order to determine the effect of post- 
mortem shrinkage and to enable these measurements to be related to the dimensions 
of the living fish, a number of specimens of C. acus were measured immediately after 
being killed in 25% alcohol and again after eighteen months preservation in 70% 
alcohol. The total length was found to decrease by about 6%, the length of the 
head by about 1% and its depth by about 2%. 



V. SYSTEMATIC ARRANGEMENT 

Family Carapidae 

(Fierasferidae of older literature and originally united with the Ophidiidae, 
commonly known as fierasfers, pearl-fish or glass-eels) . 

Familial characters. Small percomorph fishes with slender, elongate, com- 
pressed or cylindrical, scaleless bodies, tapering to an acuminate tail ; head short, 
usually thickest and deepest part of body ; snout blunt and rounded ; eye large, 
slightly oval, placed high on side of head ; posterior nostril a crescentic slit just in 
front of the orbit ; anterior nostril circular, on small papilla at about midlength 
of snout ; mouth large, usually oblique, lower jaw included within upper ; maxilla 
reaching to or beyond posterior margin of orbit ; teeth on premaxillae, dentaries, 
palatines and vomer, those of upper jaw being the smallest and those of the vomer 
usually the largest ; tongue smooth, pointed, free at tip ; gill-openings wide, the 
four membranes little united and leaving most of the isthmus uncovered ; 
branchiostegals 6 or 7 ; top posterior margin of operculum prolonged as small point 
projecting back above base of pectoral fins ; anus far forward in adult ; median 
fins long and low, inserted just behind head and continuous round tip of tail ; no 
caudal fin ; no pelvic fin or girdle ; pectoral girdle always present, but pectoral 
fin may be reduced or even absent ; sexes not distinguishable on external characters ; 
eggs pelagic ; life history complex, probably always involving a planktonic larva 
(vexillif er) and a benthic larva (tenuis) . 



Key to Genera and Subgenera — Adults only 

1 Teeth in bands on jaws and palatines ; maxilla not concealed by skin . . 2 
Teeth in single rows on jaws and palatines ; maxilla concealed by skin . . 3 

2 (1) No diastema in lower jaw ; anus almost vertically below base of pectoral . . 4 

Diastema in lower jaw ; anus clearly posterior to base of pectoral . . Echiodon 

3 (1) Pectoral fins present ....... Encheliophis (Jordanicus) 

Pectoral fins absent ....... Encheliophis (Encheliophis) 

4 (2) Anterior teeth not fang-like, body slightly compressed or cylindrical, deepest part 

usually the head ........ Carapus (Carapus) 

Anterior pair of teeth in each jaw large, fang-like ; body strongly compressed, 

deepest part behind head ....... Carapus (Onuxodon) 



26o SYSTEMATIC REVISION OF THE FAMILY TELEOST CARAPIDAE 

Genus CARAPUS Refinesque 1810 

Type Gymnotus acus Linnaeus 

Carapus Rafinesque-Schmaltz, 1810, Indice d'lttiologia Siciliana : 57. No type stated ; Gym- 
notus acus Linnaeus was designated type by Opinion 42 of the International Commission for 

Zoological Nomenclature, 191 2. 
Fierasfer Oken, 1817, Isis, 1817:1182. Name derived from " les fierasfers ", Cuvier 1815, 

Mem. Mus. Hist. nat. Paris, 1 : 119. Type Ophidium imberbe Cuvier, 1815, (non Ophidion 

imberbis Linnaeus, 1758, Sy sterna Naturae, 1 : 259). 
Diaphasia Lowe, 1843, Proc. zool. Soc, Lond. 11 : 92. Type Gymnotus acus Bninnich, 1768, 

Ichthyologia Massiliensis : 13. 
Oxybeles Richardson, 1844, Ichthyology of the Voyage of H. M.S. " Erebus " and " Terror " : 73. 

Type Oxybeles Homei Richardson, 1844. 
Porobronchus Kaup, i860, Ann. Mag. nat. Hist. (3) 6 : 272. Type Porobronchus linearis Kaup, 

i860. 
Helminthodes Gill, 1864, Proc. Acad. nat. Sci. Philadelphia, 16 : 203 (non Helminthodes Marsh, 

1864, Amer. J. Sci. 38 : 415). Type Oxybelus lumbricoides Bleeker, 1854, Nat. Tijdschr. Ned.- 

Ind. 7 : 163. ' 
Vexillifer Gasco, 1870, Bull. Assoc. Nat. Med. Napoli, 1870 : 59. Type Vexillifer dephilippii 

Gasco, 1870. 
Helminthostoma Cocco. MS name cited by Giinther, 1870, Catalogue of Fishes, 8 : 145. Type 

Helmintho stoma delle Chiaje Cocco. 
Lefroyia Jones (J. M.), 1874, Zoologist, (2) 9 : 3837. Type Lefroyia bermudensis Jones, 1874. 
Rhizoiketicus Vaillant, 1893, C.R. Acad. Sci., Paris, 117 : 745. Type Rhizoiketicus carolinensis 

Vaillant, 1893. 
Leptofierasfer Meek & Hildebrand, 1928, Publ. Field Mus. zool. Ser. 15 : 963. Type Lepto- 

fierasfer macrurus Meek & Hildebrand, 1928. 
Pirellinus Whitley, 1928, Rec. Austral. Mus. 16 : 211. Type Oxybeles lumbricoides Bleeker, 

1854. 
Disparichthys Herre, 1935, Publ. Field Mus. zool. Ser. 18 : 383. Type Disparichthys fluviatilis 
Herre, 1935. 

Generic characters. Body compressed or cylindrical ; lateral processes of 
first and second vertebrae long and not expanded, of third and fourth vertebrae 
expanded and fused together, of fifth and subsequent vertebrae short and not 
expanded ; trunk vertebrae number 17-20 ; lower jaw stout, nearly straight, 
tooth-bearing portion tapering to tip ; in adult, teeth of jaws and palatines arranged 
in bands ; distance between teeth approximately equal to diameter of tooth-base ; 
no diastema in lower jaw ; maxilla extending beyond posterior edge of orbit in 
adult, clearly outlined by folds of skin ; interorbital domed ; upper edge of orbit 
not impinging on dorsal profile of head ; anus in adult close to roots of pectoral 
fins ; swim-bladder short ; no pyloric caecae ; branchiostegals 7. 

The most familiar name for members of the Carapidae is " fierasfer ", a term first 
used by Briinnich (1768) who gave it as a local name for the species now known as 
Carapus acus in use by the fishermen of Marseilles. The name next appears as " les 
fierasfers " (Cuvier, 1815 and 1817) when it was used for a group of subgeneric rank 
under Ophidium, containing Ophidium imberbe (a synonym of Carapus acus) and 
Ophidium dentatum (now Echiodon dentatus) . This was quickly given nomenclatural 
status as Fierasfer Oken (1817), the name which has been and is still the most widely 
used for members of this family. 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 261 

However prior to Cuvier, the heterogenous Linnean genus Gymnotus had been 
divided into Gymnotus, sensu strictu, and Carapus by Rafinesque-Schmaltz (1810). 
Rafinesque did not specify a type for the new genus, nor even list the species it was 
created to contain, but gave only the following diagnosis : — 

" XII. Nessun'ala dorsale, ne caudale, un'ala anale e due pettorali, mascella 
superiore piu lunga dall'inferiore, coda nuda al disotto. Osserv. Differisce 
dal vero genere Gymnotus, che ha Fala anale lunghissima, ricuoprendo il disotto 
della coda, e la mascella inferiore piu lunga dalla superiore.' ' (XII. Neither 
dorsal nor caudal fins ; an anal fin and two pectorals ; upper jaw longer than 
the lower ; tail naked below. Notes. It differs from the true genus Gymnotus, 
which has the anal fin very long, covering the lower part of the tail, and the 
lower jaw longer than the upper.) 

In another part of his account, Rafinesque cited Gymnotus acus Linnaeus as a 
Sicilian representative of Carapus. By Opinion 42, rendered 191 2, the International 
Commission for Zoological Nomenclature has designated Gymnotus acus Linnaeus 
(no date given) as type of Carapus Rafinesque, invalidating Fierasfer as a generic name. 

Gymnotus acus Linnaeus does not, strictly, exist at all and the characters used by 
Rafinesque in his diagnosis of Carapus are such as to exclude from this genus all 
species of fierasfer. Rafinesque derived his nomenclature from the 13th edition of 
the Sy sterna Naturae, edited by Gmelin (1788), in which is a shortened description 
of Gymnotus acus Brunnich, correctly attributed, the species that Rafinesque 
presumably intended to name. The diagnosis of Carapus is apparently a translation 
into Italian of Gmelin's Latin description of the South American fresh-water fish 
Gymnotus carapo Linnaeus, type species of the genus Gymnotus Linnaeus. The 
resemblance of the new generic name to the older trivial name emphasizes the 
similarity of the descriptions. 

Of the remaining generic synonyms, Diaphasia Lowe was proposed as an alterna- 
tive to the then current name Fierasfer. The species described by Richardson and 
Jones are so closely akin to Carapus acus that generic separation of either is un- 
warranted, while Rhizoiketicus Vaillant must be united with Carapus for lack of 
distinguishing characters. This genus was created to contain a fierasfer from the 
Caroline Islands, the two examples of which were described as having large, easily- 
detachable, lozenge-shaped scales above and below the lateral line. These were 
not true scales, but were formed from and continuous with the outer cornified 
layers of the skin. Desiccation of any long-preserved fierasfer will cause cracking 
and flaking of the skin and these " scales " are thus no adequate reason for retaining 
Rhizoiketicus as a separate genus. The other characters of the two specimens are 
not recorded and they seem to be no longer in existence, nor has the species been 
found since. The other generic synonyms are based upon larvae. Porobronchus 
Kaup and Vexillifer Gasco belong to C. acus, Helminthodes Gill, and its substitute 
Pirellinus Whitley, probably to C. homei, and Leptofierasfer Meek & Hildebrand 
probably to C. dubius ; it seems likely that Disparichthys Herre may ultimately 
also be found to be attributable to a species of Carapus. 

Recently, Carapus parvibrachium Fowler has been made the type of a new genus, 

zool. 4, 6. 18 



262 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

Onuxodon Smith, 1955, on the grounds that it differs from all other fierasfers in 
possessing an extremely deep, strongly compressed body, strongly domed inter- 
orbital and fewer vertebrae. To these characters may be added a partly calcified 
swim-bladder and a slightly greater number of trunk vertebrae than occurs in most 
species of Carapus. In all these features the species originally described as 
Fierasfer margaritiferae Rendahl agrees with C. parvibrachium and there can be no 
doubt that the two are congeneric. However, in other respects — vertebral and 
jaw structure, position of the anus, dentition, general body proportions — these two 
species are more closely akin to Carapus spp. than to the other fierasfers and for this 
reason it is considered that their probable evolutionary relationships are more 
truly expressed by regarding C. parvibrachium and F. margaritiferae as representing 
a subgenus of Carapus. In the following account, therefore, the genus is divided 
into two subgenera, Carapus and Onuxodon. It is not improbable that future 
investigations will result in the creation of a third subgenus to contain Carapus 
parvipinnis (Kaup) , but this course is not at present justified. 

Subgenus CARAPUS 
Subgeneric characters. Body slightly compressed or cylindrical, not so deep 
as length of head ; interorbital flat or only slightly domed, its width equal to or 
greater than horizontal diameter of eye ; trunk vertebrae 17-18; swim-bladder 
uncalcified. 

Key to Species — Adults only 

1 Head more than one-ninth of total length ; pectoral fins more than one-third 

length of head ............ 2 

Head less than one-ninth of total length ; pectoral fins less than one-third length 

of head ........... C. parvipinnis 

C. boraborensis 

2 (1) No enlarged teeth at front of jaws ........ 6 

Enlarged teeth at front of upper jaw ........ 3 

Enlarged teeth at front of upper and lower jaws ...... 9 

3 (2) Median row of large teeth on vomer, flanked by small teeth .... 4 

Anterior vomerine teeth small ; posterior vomerine teeth large . . C. birpex 

4 (3) Teeth of upper jaw uniserial ......... C. pindae 

Teeth of upper jaw polyserial ......... 5 

5 (4) Outermost series of teeth in lower jaw considerably larger than inner series . C. homei 

Outermost series of teeth in lower jaw not conspicuously larger than inner series 

C. dubius 

C. bermudensis 

6 (2) Maximum depth of body considerably greater than maximum depth of head . 8 

Maximum depth of body not greater than maximum depth of head ... 7 

7 (6) Anus anterior to roots of pectoral fins ....... C. acus 

Anus vertically beneath roots of pectoral fins . . . . . C. kagoshimanus 

8 (6) Pectoral one-third length of head ; body progressively tapering from region of 

maximum depth to tip of tail . . . . . . . C. houlti 

Pectoral more than one-half length of head ; body narrowing sharply at end of 

abdomen before tapering to tail tip . . . . . . . C. cuspis 

9 (2) Head one-eighth to one-ninth of total length (about 200 mm.) ; outer series of 

teeth in lower jaw similar to inner series ..... C. owasianus 

Head one-sixth to one-seventh of total length (about 100 mm.) ; outer series of 

teeth in lower jaw much taller than inner series . . . . . C. caninus 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 263 

Carapus acus (Briinnich), 1768 
(Text-fig. 6) 

Gymnotus acus Briinnich, 1768, Ichthyologia Massiliensis : 13. Artedi, 1788, Bibliotheca Ich- 

thyologica : 164. Gmelin, 1788, Sy sterna Naturae Linnaei, 1 : 1140. 
Notopterus Fontanesii Risso, 1810, Ichthyologie de Nice : 82. 
Ophidium imberbe Cuvier, 1815, Mem. Mus. Hist. Nat., Paris, 1 : 119. (non Ophidion imberbis 

Linnaeus, 1758, Sy sterna Naturae, 1 : 259.) Cuvier, 181 7, Regne Animal, 2 : 259. 
Fierasfer imberbe, Oken, 1817, Isis, 1817 : 1182. 
Ophidium fierasfer Risso, 1826, Histoire Naturelle des Principales Productions de V Europe Meri- 

dionale, 3 : 212. 
Fierasfer fontanesii, Costa (O.), 1829, Fauna del Regno di Napoli, 3 : tab. 20 bis. 
Ophidium sp., Delle Chiaje, 1841, Descrizione e Notomia degli Animali Invertebrati della Sicilia 

Citeriore, 4:3. 
Diaphasia acus, Lowe, 1843, Proc. zool. Soc, Lond. 11 : 92. 
Fierasfer acus, Kaup, 1856, Arch. Naturges. 22 : 93. Kaup, 1856, Catalogue of Apodal Fish : 157. 

Giinther, 1862, Catalogue of Fishes, 4 : 381. Steindachner, 1868, S.B. Akad. Wiss. Wien, 

57 : 46. Canestrini, 1872, Fauna d'ltalia : Pesci : 191. Emery, 1878, Atti. Soc. ital. Sci. 

nat., Milano, 21 : 37. Emery, 1880, Fauna u. Flora Neapel, 2 : 1. Giglioli, 1880, Esposi- 

zione di Pesca : 97. Perugia, 1881, Elenco dei Pesci delV Adriatica : 38. Emery, 1882, Mitt. 

zool. Stat. Neapel, 3 : 281. Carus, 1885, Prodromus Faunae Mediterranae, 2 : 580. Raffaele, 

1888, Mitt. zool. Stat. Neapel, 8 : 39. Lo Bianco, 1904, Pelagische Tiefseefischerei der Maja 

in der Umbegung von Capri : 24. Soljan, 1948, Ribe Jadrana : 122. 
Helmintho stoma delle Chiaje Cocco. MS name cited by Giinther, 1870, Catalogue of Fishes, 

8 : 145. 

Fierasfer massiliensis Briinnich. Name cited by Kaup, 1856, Catalogue of Apodal Fish : 157, 

without reference. 
Porobronchus linearis Kaup, i860, Ann. Mag. nat. Hist. (3) 6 : 272. Giinther, 1870, 

Catalogue of Fishes, 9 : 145. Belotti, 1891, Atti Soc. ital. Sic. nat., Milano, 33 : 127. 
Vexillifer dephilippii Gasco, 1870, Bull. Assoc. Nat. Med., Napoli, 1870 : 59. 
Vexillifer de Filippii, Costa (A.), 1871, Ann. Mus. zool., Napoli, 6 : 88. 
Encheliophis tenuis Putnam, 1874, Proc. Boston Soc, nat. Hist. 16 : 343. 
Fierasfer dentatus (part), Emery, 1880, Fauna u. Flora Neapel, 2 : 16. 
Fierasfer dentatus, Emery, 1882, Mitt. zool. Stat. Neapel, 3 : 283. 
Fierasfer imberbis, Moreau, 1881, Histoire Naturelle des Poissons de la France, 3 : 226. Moreau, 

1892, Manuel d' Ichthyologie Francaise : 405. 
Carapus imberbis, Fowler, 1936, Bull. Amer. Mus. nat. Hist. 70 : 1073. 
Carapus acus, Padoa, 1947, Pubbl. Staz. zool. Napoli, 20 : 111. Arnold, 1953, Pubbl. Staz. zool. 

Napoli, 24 : 153. 

The material examined includes 39 specimens in the collection of the British 
Museum (Natural History) ; 17 in the collection of the Institut Oceanographique de 
Monaco ; 22 in the collection of the Universitetets Zoologiske Museum, Copenhagen ; 
40 in the Naples Sales Collection, 1952 ; and 29 in the Naples Sales Collection, 1954. 
Of these 147 specimens the majority were adults or juveniles, only a few being 
vexillifer or tenuis larvae. The localities represented were Sicily, Naples, Genoa, 
Monaco and Nice. The type, which was obtained at Marseilles, France, may be no 
longer in existence. 

Specific characters. Adult : Maximum length about 200 mm. ; length of head 
one-seventh to one-eighth of total length ; maximum depth of head one-half and 
maximum width one-third of length of head ; horizontal diameter of eye equal to 
length of snout and to interorbital width ; mouth very oblique ; maxilla extends 



264 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

only a short distance posterior to orbit ; jaw teeth small and uniform ; vomerine 
teeth a little longer than jaw teeth, arranged in three slightly irregular, longitudinal 
rows, those of the central row being the largest ; pectoral fins one-half length of 
head ; anus anterior to roots of pectorals ; body slightly translucent in life, heavily 
blotched or barred with red ; about fifteen golden or silvery spots along flanks just 
above lateral line ; iris silver ; operculum and abdomen often with silvery lustre. 

Juvenile : Adult colouration first assumed at about ioo mm. length ; general 
body form resembles that of adult, though head of younger individuals may be but 
one-ninth or one-tenth of total length ; dentition and proportions of head similar 
to those of adult ; maxilla extends little beyond hind edge of orbit ; pectorals one- 
third or less of length of head ; anus beneath or slightly anterior to roots of 
pectorals ; body translucent, spotted or barred with dark-brown or black ; no sign 
of red pigment whatsoever ; golden or silvery spots along sides as in adult. 

Tenuis larva : Maximum recorded length more than 200 mm.; body slender and 
cylindrical, not compressed as in juvenile and adult ; head less than one-twelfth 
of total length ; maxilla extends only as far as middle of orbit ; teeth in single rows 
on jaws and palatines ; enlarged teeth may be present at front of one or both jaws ; 
pectoral fins less than one-fifth length of head ; anus posterior to roots of pectorals ; 
transparent and unpigmented in life. 

Vexillifer larva : Maximum recorded length more than 80 mm.; head about one- 
tenth of total length ; maxilla extends only to anterior edge of orbit ; teeth in 
single rows on jaws and palatines ; older vexillifers have enlarged teeth at front of 
one or both jaws ; pectoral fins about one-eighth of length of head ; anus near 
posterior end of abdomen ; body translucent in life and devoid of pigmentation. 

Eggs : Spawning period July-September ; eggs adhere in oval, yellowish rafts 
about 80 mm. in length. 

The main dimensions of the adult fish studied are summarized in Table I and 
measurements and proportions of representative specimens are given in Table II. 



Table I. — Carapus acus (Brilnnich,) 1768. Summary of 
Measurements and Proportions of Adults. 

R. M ± a M . 

Variate. N. mm. mm. 

Total length . . . 72 

Length of head . . . 73 

Depth of head . . . 72 

Width of head ... 49 



Length of pectoral . . 49 

Preanal length . . .49 



Length of head (% TL) . 72 

Depth of head (% HL) . 72 

Width of head (% HL) . 53 

Length of pectoral (% HL) 48 

Preanal length (% TL) . 47 



100-202 

13-28 

7-16 

4-1 1 

7-16 

13-23 



% 
11.6-15.3 
43.8-71-4 
26-7-50.0 
38-8-64-0 
10-8-13-3 



i55*4±2-34 
20'6±i *o8 
ii-3±o-23 

8.0^0*25 
10-6^0-30 

18.7io.37 



i3*2±o-o8 
55-o±o-69 
38-6^0-23 
51.6io.84 
11 .6^0-10 



N = number of specimens ; R = range of variate ; M = mean of variate (± standard error of 
mean, a M ) ; TL = total length ; HL = length of head. 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 265 

Table II. — Carapus acus (Brunnich), 1768. Measurements 
and proportions of representative adults. 

Collection .... 

Museum number 

Locality .... 



Total length 
Length of head . 

Maximum depth of head . 

Maximum width of head . 

Length of snout 

Horizontal diameter of eye 

Verical diameter of eye . 

Interorbital width . 

Length of maxilla . 

Length of pectoral fin 
Preanal length . 
Maximum depth of body 

TL = to 

C. acus is quite common in the western Mediterranean, being recorded from the 
coastal waters of Spain, France, Italy, Sicily, Sardinia and the Balearic Islands. It 
may also occur in the Adriatic, Aegean and off the Algerian coast and has occasionally 
been reported in the Atlantic. These latter records, however, are of doubtful 
validity. C. acus has occasionally been taken free-swimming, but all stages except 
the vexillifer normally live within the body cavities of holothurians (Holothuria 
tubulosa and Stichopus regalis), where they attack the gonads and branchial trees. 
The tenuis larva is entirely dependent upon its host and does not survive long in 
sea water. 



British Museum 


1952. 11. 25. 1 




1952. 11. 25. 5 


• 


Naples 




mm. 




mm. 


184 




152 


24 (13% TL) 






20 (13% TL) 


14 (58% HL) 






11 (55% HL) 


10 (42% HL) 






9 (45% HL) 


4 (17% HL) 






3-5 (i 7 '5% HL) 


4 (17% HL) 






3'5 (i 7 -5% HL) 


3-5(i5%HL) 






3-2 (16% HL) 


4 (17% HL) 






4 (20% HL) 


11 (46% HL) 






9-5 (47'5% HL) 


12 (50% HL) 






10 (50% HL) 


22 (12% TL) 






18 (12% TL) 


14.5 (60% HL) 






i3'5 (67-5% HL) 


length ; HL = length of head. 







Carapus birpex n. sp. 

(Text-fig. 7) 

The type, the only specimen known, is an adult, found in a holothurian taken 
at Madeira, precise locality unknown. It is in the collection of the Museu Municipal, 
Funchal, Madeira, museum no. 2739. Two other fierasfers were found in the same 
holothurian, but it is unknown whether they have been preserved. 

Specific characters. Length of type 209 mm. ; head one-seventh of total length ; 
maximum depth of head more than three-fifths and maximum width one-fifth of 
length of head ; horizontal diameter of eye equal to length of snout and slightly 
less than interorbital width ; mouth very oblique ; maxilla extends behind orbit 
for distance equal to half horizontal diameter of eye ; owing to obliquity of mouth, 
end of maxilla almost cuts ventral profile of lower jaw ; single pair of enlarged teeth 
at front of upper jaw, the rest, like those of the lower jaw, small and uniform ; 
anterior half of vomer bears two slightly irregular rows of small, sharp, conical 
teeth, at either side of which the antero-lateral surfaces of vomer are smooth and 
toothless ; posterior portion of vomer has a single median row of five, large, stout, 



266 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 





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f 



'X 









f 



'.****£. 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 267 

curved and sharply-pointed teeth, flanked by a number of smaller teeth similar to 
those on front of vomer ; pectoral fins about two-fifths length of head ; anus 
anterior to roots of pectorals ; colour in life unknown. The other stages in the life 
history are unidentified. 

The dimensions of the type are given in Table III. 

Table III. — Carapus birpex n. sp. Measurements and Proportions of the Type. 



Museum .... 


Museu Municipal, Funchal 


Locality .... 


Madeira, 
mm. 




Total length .... 


209 




Length of head 


29 


(14% TL) 


Maximum depth of head 


15-5 


(58% HL) 


Maximum width of head 


11 


(39% HL) 


Length of snout 


5*5 


(19% HL) 


Horizontal diameter of eye 


5'5 


(19% HL) 


Vertical diameter of eye 


5 


(17% HL) 


Interorbital width 


6 


(21% HL) 


Length of maxilla 


15 


(52% HL) 


Length of pectoral fin 


12 


(4i% HL) 


Maximum depth of body 


18 


(62% HL) 


Preanal length 


25 


(12% TL) 


TL = total length ; HL = 


length of head. 


Carapus cuspis n. sp. 




(Text-fig. 8) 





The three known examples of this species were taken free-swimming near Madeira, 
position 33 02' N., 16 20' W., depth 100 metres (about 50 fathoms), during the 
1897 cruise of the yacht " Princesse- Alice ". They were recorded, without descrip- 
tion or figure, as Fierasfer acus by Roule (1917) . The smallest of the three specimens 
is designated type and this and one paratype are in the collection of the Institut 
Oceanographique de Monaco. A second paratype is in the collection of the British 
Museum (Natural History) . 

Specific characters. Length of type 216 mm. (of paratypes 221 mm. and 230 
mm.); head a little more than one-eighth of total length ; maximum depth of head 
two-thirds and maximum width one-half of length of head ; horizontal diameter 
of eye slightly greater than length of snout, but less than interorbital width ; mouth 
oblique ; maxilla extends behind orbit for distance equal to half horizontal diameter 
of eye ; teeth on jaws and palatines resemble those of C. acus ; vomer with single 
row of rather large, sharp, backwardly-directed teeth, flanked by a few smaller 
ones ; anal fin deep, fleshy anteriorly ; pectorals three-fifths length of head ; anus 
slightly anterior to roots of pectoral fins. Colour in life unknown. Other stages in 
the life history have not been identified. 

The body does not taper steadily from head to tail-tip, as in most fierasfers. 
Instead, the abdomen is deep and rounded and the maximum depth of the body is 



268 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

about 40% greater than the maximum depth of the head. Beyond the abdomen 
the body narrows sharply to the long, tapering tail. 

The paratypes closely resemble the type, differing only in their dimensions and 
in some minor details of their proportions. Through long preservation in alcohol 
their colour has been entirely destroyed, but a record from Algiers (Guichenot, 1850) 
may perhaps refer to this species. In this account a fierasfer, said to be common 
along the Algerian coast, was identified as Fierasfer imberbe and described as having 
a yellowish ground colour crossed by numerous brown bands, the abdomen bluish 
and faintly spotted with red. This colour scheme is quite unlike that characteristic 
of C. acus. 

The dimensions of the type and paratypes of C. cuspis are given in Table IV. 

Table IV. — Carapus cuspis n. sp. Measurements and 
Proportions of Type and Two Paratypes. 



Collection . 


• 


Monaco 




Monaco 


British 
Museum. 


Status 




Type 


• 


Paratypes 1 and 2 




mm. 




mm. 




mm. 


Total length 


216 




221 




230 


Length of head . 


28 


(i3-o% TL) 


30 


(13-6% TL) 


31 (i 3 -5% TL) 


Maximum depth of head 


17 


(61% HL) 


17 


(57% HL) 


18 (58% HL) 


Maximum width of head 


13 


(46% HL) 


13 


(43% HL) 


14 (45% HL) 


Length of snout 


5 


(18% HL) 


5 


(17% HL) 


5 (16% HL) 


Horizontal diameter of ey( 


1 6 


(21% HL) 


6 


(20% HL) 


6 (19% HL) 


Vertical diameter of eye 


5 


(18% HL) 


5 


(17% HL) 


5 (16% HL) 


Interorbital width . 


6- 


5 (23% HL) 


6-f 


(22% HL) 


7 (23% HL) 


Length of maxilla . 


15 


(54% HL) 


15 


(50% HL) 


17 (55% HL) 


Maximum depth of body 


24 


(86% HL) 


24 


(80% HL) 


22 (71% HL) 


Length of pectoral fin 


17 


(61% HL) 


17 


(57% HL) 


17 (65% HL) 


Preanal length . 


27 


(2-5% TL) 


27 


(13% TL) 


28 (12% TL) 




TL = total length ; HL = 


= length 


of head. 





The two new species, C. birpex and C. cuspis, are both from the same general 
locality and both bear a superficial resemblance to C. acus, to which species they are 
probably most closely related. They differ from it and from each other especially 
in their dentition, the size and form of their maxillae and the general shape and 
proportions of the head, as well as in absolute size. Of the two, C. birpex is the 
furthest removed from C. acus. The snout is considerably more pointed than is 
that of C. acus when viewed from the side, squarer when viewed from above (Text- 
fig. 9). Further, though the head of C. birpex is longer than that of the largest 
C. acus available for study, yet the snout is of no greater length. Hence the pre- 
orbital portion of the skull is relatively shorter in C. birpex, the post orbital relatively 
longer. Another important difference between the two species is the obliquity of the 
mouth, which in C. birpex is so great that the maxilla almost impinges on the ventral 
profile of the lower jaw, though it does not in fact extend so far behind the orbit as 
does the maxilla of C. acus. Finally, C. birpex is separated from C. acus by the 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 269 

possession of enlarged teeth at the front of the upper jaw and by the dual nature 
of the vomerine teeth, which in form and arrangement are quite unlike those of any 
other species of Carapus so far described. C. birpex resembles the Indo-Pacific 
species C. homei in having enlarged jaw teeth, but differs from this species in the 
absence of an outer row of strong teeth in the lower jaw. 

C. cuspis, on the other hand, resembles C. acus in many respects, particularly 
in the dentition of the jaws and the proportions of the head. But the maxilla is 
relatively shorter than is that of C. acus, though the obliquity of the mouth is no 
greater, and the median row of vomerine teeth are larger and more strongly curved. 
C. cuspis is most clearly distinguished by the depth and rotundity of the abdomen 
and especially by the constriction of the body at the base of the tail. 






B 



20 mm 

Fig. 9. — Heads of European Carapus spp. from the dorsal aspect. 
a. C. acus ; b, C. birpex ; c, C. cuspis. 



C. acus has been recorded from Atlantic localities upon a number of occasions and 
under a variety of names : 

Madeira, as Diaphasia acus (Lowe, 1843). 
Canary Islands, as Fierasfer acus (Vinciguerra, 1893). 
Portuguese coast, as Fierasfer imberbis (Nobre, 1935). 
Portuguese coast, as Carapus imberbis (de Buen, 1935). 
West Africa, as Carapus imberbis (Fowler, 1936). 
Cape Verde Islands, as Carapus imberbis (Cadenat, 1937). 
Biarritz, as Fierasfer imberbis (Pellegrin, 1937). 
Senegal, as Carapus imberbis (Cadenat, 1950). 



270 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

Of these authors, two alone (Nobre, 1935 ; Fowler, 1936) provided a description, 
both of which were based upon Mediterranean specimens, not on those from the 
locality at which the species was reported. Nobre included a figure, but this is 
unrecognizable. It seems probable that these records relate either to C. birpex or 
to C. cuspis, not to C. acus at all. 



Carapus dubius (Putnam), 1874 
(Text-fig. 10 a) 

Fierasfer dubius (part) Putnam, 1874, Proc. Boston Soc. nat. Hist. 16 : 339. Jordan & Gilbert, 

1882, Bull. U.S. nat. Mus. 16 : 791. 
Fierasfer arenicola Jordan & Gilbert, 1881, Proc. U.S. nat. Mus., 4 : 338. Jordan, 1895, Proc. 

Calif. Acad. Sci. (2) 5 : 502. Jordan & Evermann, 1898, Bull. U.S. nat. Mus. 47 : 2183. 
Fierasfer affnis, Jordan & Evermann, 1898, Bull. U.S. nat. Mus., 47 : 2183. (non Giinther, 

1862, Catalogue of Fishes, 4 : 381.) 
Carapus dubius, Meek & Hildebrand, 1928, Publ. Field Mus. zool. Ser. 15 : 963. 
Leptofierasfer macrurus Meek & Hildebrand, 1928, Publ. Field Mus. zool. Ser. 15 : 963. 
Carapus affinis (part), Rivero, 1936, Proc. Boston Soc. nat. Hist. 41 : 41. (non Giinther, 1862.) 




25 mm 



2 mm 




25 mm 



B 



Fig. 10. — a, Carapus dubius (Putnam), and its vomerine teeth, b, Carapus bermudensis 

(Jones), and its vomerine teeth. 



The following account is based upon a single adult in the collection of the British 
Museum (Natural History) from the Gulf of Panama. 

Putnam did not designate a type. His description was based upon fourteen 
specimens, eight from Pearl Island, in the Gulf of Panama, the rest from Florida, 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 271 

the Bahamas and the Tortugas. Most of these specimens, including five from 
Panama, are in the Museum of Comparative Zoology, Harvard. In Putnam's list 
of specimens those from Pearl Island stand first ; they must therefore be regarded as 
the cotypes and Pearl Island, Gulf of Panama, as the type locality. 

Specific characters. Maximum length apparently about 100 mm.; head one- 
seventh to one-eighth of total length ; maximum depth of head one-half to three- 
fifths and maximum width one-third to two-fifths of length of head ; horizontal 
diameter of eye equal to length of snout, greater than interorbital width ; mouth 
slightly oblique ; maxilla extending only a short distance behind posterior edge of 
orbit ; teeth of upper jaw small, one or two anterior ones enlarged ; outermost 
series of lower jaw slightly stouter than inner series ; row of about four large, stout, 
conical teeth on vomer, flanked by smaller ones ; pectoral fin three-fifths length of 
head ; anus anterior to roots of pectorals ; translucent in life, a few yellowish spots 
on sides of body, tip of tail dusky (Jordan & Gilbert, 1881). Vexillifer larvae 
attributable to this species have been described under the name Leptofierasfer 
macrurus Meek & Hildebrand (1928). 

The measurements and proportions of the only specimen available for study are 
given in Table V. 

Table V. — Carapus dubius (Putnam), 1874, and C. bermudensis (Jones), 1874. 
Measurements and Proportions of Adult Specimens. 



Collection 


British Museum 


Species 


C. dubius 


C. bermudensis 


Museum number . 


I935-3-24. 26 


1931.9.24.1 


Locality- 


Gulf of Panama 


Florida 




mm. 


mm. 


Total length 


108 


104 


Length of head 


16 (15% TL) 


15 (14% TL) 


Maximum depth of head 


10 (62-5% HL) 


7 (47% HL) 


Maximum width of head 


7 (44% HL) 


5 (33% HL) 


Length of snout 


3-5 (22% HL) 


2.5 (17% HL) 


Horizontal diameter of eye 


3.5 (22% HL) 


2-5 (17% HL) 


Vertical diameter of eye 


3-2 (20% HL) 


2 (13% HL) 


Interorbital width 


3 (19% HL) 


3 (20% HL) 


Length of maxilla 


8 (50% HL) 


6-5 (43% HL) 


Length of pectoral fin . 


10 (62-5% HL) 


7 (47% HL) 


Maximum depth of body 


11 (69% HL) 


7.5 (50% HL) 


Preanal length 


14 (13% TL) 


12 (12% TL) 


TL = tota 


1 length ; HL = length of hea 


a. 



C. dubius has been recorded from the Pacific coast of Central America, including 
Pearl Island and Chame Point, Gulf of Panama ; Mazatlan, Mexico ; and the Gulf 
of California. Examples have been found lying free in the sand, but the majority 
of specimens have been taken as inquilines of lamellibranchs. 

The taxonomic status of the fierasfers from the Pacific and Atlantic sides of 
Central America is a matter concerning which, through lack of specimens, no final 



272 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

decision can yet be made. In 1874, two species, both undoubted Carapus, were 
described — Lefroyia bermudensis Jones from Bermuda and Fierasfer dubius Putnam 
from the Gulf of Panama and West Indies. Neither was figured, the type of L. 
bermudensis cannot be traced, and the original descriptions are distinguishable 
neither from each other nor from that of the Indo-Pacific species C. homei. Most 
authors have regarded C. dubius and C. bermudensis as conspecific and many have 
further equated them with Fierasfer affinis Gunther. Further complications were 
introduced by the designation of Fierasfer arenicola Jordan & Gilbert and Lepto- 
fierasfer macrurus Meek & Hildebrand, a vexillifer larva. Fierasfer arenicola is 
now generally accepted as a synonym of F. dubius, and the larval nature of L. 
macrurus has also been recognized (Parr, 1930). 

Now, it is quite clear that all these forms, whatever their status, belong to the one 
genus Carapus. Further, there is no reason for uniting F. affinis, the locality of 
which is unknown, with the American fierasfers. This species, the type of which 
is in the British Museum, is far larger than any described from American water and 
(p. zyy) is not distinguishable from C. homei. 

Putnam based his description of F. dubius upon specimens from the Gulf of 
Panama, but was unable to distinguish them from others from the West Indies. 
The identity of the two forms is supported by Rivero (1936) who re-examined such of 
Putnam's specimens as are still available and compared them with other material 
from Cuba and Jamaica. It appears best, however, to consider these Pacific and 
Atlantic fierasfers as two closely-related species, pending re-examination of the 
problem on the basis of more adequate material. The trivial name dubius should be 
restricted to the Pacific fierasfers, while the trivial name bermudensis is available for 
those from the Atlantic. 

Carapus bermudensis (Jones), 1874 
(Text-fig. iob) 

Lefroyia bermudensis Jones (J. M.), 1874, Zoologist (2) 9 : 3837. 

Fierasfer dubius (part), Putnam, 1874, Proc. Boston Soc. Nat. Hist. 16 : 343. Jordan & Gilbert, 

1882, Bull. U.S. nat. Mus. 16 : 791. 
Fierasfer bermudensis, Jordan & Evermann, 1898, Bull. U.S. nat. Mus., 47 : 2443. Herre, 

1942, Stanford Univ. Publ. biol. Sci. 7 (2) : 20. 
Fierasfer dubius, Parker, 1926, Proc. nat. Acad. Sci. Washington, 12 : 421. 
Carapus dubius (part), Meek & Hildebrand, 1928, Publ. Field Mus. zool. Ser. 15 : 963. 
Fierasfer affinis, Breder, 1929, Field Book of Marine Fishes of the Atlantic Coast : 279. (non 

Gunther, 1862.) 
Carapus affinis, Rivero, 1936, Proc. Boston. Soc. nat. Hist. 41 : 41. (non Gunther, 1862.) 
Carapus sp., juv., Parr, 1930, Bull. Bingham Oceanogr. Coll. 3 : 133. 
Carapus bermudensis, Longley & Hildebrand, 1941, Papers Tortugas Lab. 34 : 90. Beebe & 

Tee-Van, 1933, Field Book of the Shore Fishes of Bermuda : 232. 
Fierasfer sp., larva, Herre, 1942, Stanford Univ. Publ. biol. Sci. 7 (2) : 20. 

The specimens examined are five adults and six larvae in the collection of the 
British Museum (Natural History). The localities represented are (adults) Florida, 
Cayman Islands, St. Kitts, Antigua, (larvae) Grenada, Tobago, Antigua. The 
type was obtained at Bermuda, but is probably no longer in existence. 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 273 

Specific characters. Maximum length probably less than 150 mm ; head 
one-seventh to one-eighth of total length ; maximum depth of head one-half and 
maximum width two-fifths of length of head ; horizontal diameter of eye greater 
than length of snout and interorbital width ; mouth oblique ; maxilla not extending 
far behind posterior edge of orbit ; outermost teeth of lower jaw slightly larger than 
those of inner series ; two to four large conical teeth on vomer, flanked by smaller 
teeth ; in upper jaw, anterior teeth enlarged ; according to Rivero (1936) there is a 
single row of small, sharp, backwardly-directed teeth along the outer side of the 
premaxilla, hidden by the lips, but these are not present in all specimens ; pectoral 
fins two-fifths to one-half length of head ; anus anterior to roots of pectoral fins ; 
translucent in life, with irregular transverse reddish bands on trunk and silvery bar 
along sides (Longley & Hildebrand, 1941). There is a vexillifer larva attributable 
to this species. 

Measurements and proportions of a representative specimen are given in Table V. 

C. bermudensis occurs in the west central Atlantic area, including Bermuda, Florida, 
the Bahamas, Cuba, Jamaica, Cayman Islands, Haiti, Leeward Islands, Windward 
Islands, Trinidad and Tobago. It is inquiline in the holothurian Actinopyga agassizi. 



Carapus hornet (Richardson), 1844 
(Text-fig. n) 

Oxybeles Homei Richardson, 1844, Ichthyology of the Voyage of H. M.S. Erebus and Terror : 73. 

Bleeker, 1855, Verh. Akad. Amsterdam, 2 : 15. 
Oxybelus Brandesii Bleeker, 1851, Nat. Tijdschr. Ned.-Ind. 1 : 278. Bleeker, 1851, Nat. Tijdschr. 

Ned.-Ind. 2 : 228. Bleeker, 1852, Nat. Tijdschr. Ned.-Ind. 3 : 238. Bleeker, 1852, Verh. 

Bat. Gen. 24 : 24. Bleeker, 1854, Nat. Tijdschr. Ned.-Ind. 7 : 162. Bleeker, 1854, Nat. 

Tijdschr. Ned.-Ind. 7 : 495. Bleeker, 1858, Nat. Tijdschr. Ned.-Ind. 15 : 255. 
? Oxybelus lumbricoides Bleeker, 1854, Nat. Tijdschr. Ned.-Ind. 7 : 163. 
Fierasfer neglectus Peters, 1855, Arch. Naturges. 21 : 260. Giinther, 1862, Catalogue of Fishes, 

4 : 381. Regan, 1908, Trans, linn. Soc. Lond. 12 : 220. Barnard, 1927, Ann. S. Afr. Mus. 

21 : 884. 
Fierasfer Homei, Kaup, 1856, Catalogue of Apodal Fish : 157. Putnam, 1874, Proc. Boston Soc. 

nat. Hist. 16 : 343. Bleeker, 1863, Ned. Tijdschr. Dierk. 1 : 236. Bleeker, 1863, Ned. 

Tijdschr. Dierk. 1 : 272. Bleeker, 1865, Ned. Tijdschr. Dierk. 2 : 293. Alleyne & Macleay, 

1876, Proc. linn. Soc, N.S.W. 1 : 347. 
Fierasfer Brandesii, Bleeker, 1858, Nat. Tijdschr., Ned.-Ind. 15 : 204. Bleeker, 1858, Nat. 

Tijdschr. Ned.-Ind. 15 : 461. 
Fierasfer homei, Giinther, 1862, Catalogue of Fishes, 4 : 381. Day, 1889, The Fauna of British 

India, Fishes, 2 : 436. Fowler, 1900, Proc. Acad. nat. Sci. Philadelphia, 1900 : 523. John- 
stone, 1904, Ceylon Pearl Oyster Fisheries, 1904, Suppl. Rep. no. 15 : 211. Jordan & Ever- 

man, 1903, Bull. U.S. Fish Comm. 23 : 505. Jordan & Seale, 1905, Bull. Bur. Fish. 25 : 435. 

Jordan & Seale, 1906, Bull. Bur. Fish. 26 : 48. Jordan & Snyder, 1906, Bull. Bur. Fish. 26 : 

217. Regan, 1908, Trans, linn. Soc. Lond. 12 : 220. 
Fierasfer affinis Giinther, 1862, Catalogue of Fishes, 4 : 381. 
? Fierasfer lumbricoides, Bleeker, 1865, Ned. Tijdschr. Dierk. 2 : 192. Regan, 1908, Trans, linn. 

Soc. Lond. 12 : 220. 
? Helminthodes lumbricoides, Gill, 1864, Proc. Acad. nat. Sci. Philadelphia, 16 : 203. 
Fierasfer acus, Bleeker, 1879, Verh. Akad. Amsterdam, 18 : 21. 
Fierasfer homii, Waite, 1897, Mem. Austral. Mus. 3 : 194. 



274 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

Fierasfer microdon Gilbert, 1905, Bull. U.S. Fish Comm. 23 : 655. Jordan & Seale, 1905, Bull. 

Bur. Fish. 25 : 435. 
Fierasfer homei, Steindachner, 1906, SitzBer. Akad. Wiss. Wien, 115 : 1419. 
Fierasfer Sluiteri Weber, 191 3, Siboga-Expeditie, 32 : 95. 
? Pirellinus lumbricoides , Whitley, 1928, Rec. Austral. Mus. 16 : 226. 
Carapus homei, Fowler & Bean, 1927, Proc. U.S. nat. Mus. 71 (10) : 15. Fowler, 1928, Mem. 

Bishop Mus. 10 : 445. Mukerji, 1932, Rec. Indian Mus. 34 : 567. Herre, 1936, Publ. Field 

Mus. zool. Ser. 21 : 416. Herre, 1939, Rec. Indian Mus. 41 : 574. Abe, 1939, Palao trop. 

biol. Stud. 1 : 574. de Beaufort & Chapman, 1951, Fishes of the I ndo- Australian Archipelago, 

9 : 4496. Smith, J. L. B., 1955, Ann. Mag. nat. Hist. (12) 8 : 414. 
? Fierasfer arenicola, Borodin, 1930, Bull. Vanderbilt oceanogr. (Mar.) Mus. 1 : 62. 
Fierasfer Mourlani Petit, 1934, Bull. Mus. Hist. nat. Paris, 6 : 393. 
Carapus neglectus, Smith (J. L. B.), 1949, Sea Fishes of Southern Africa : 359. Smith (J. L. B.), 

1955, Ann. Mag. nat. Hist. (12) 8 : 413. 

The specimens examined include the type and 48 others (27 adults, 2 tenuis, the 
rest probably juveniles) in the collection of the British Museum. The localities 
represented are the Seychelles, Laccadives, Chagos Archipelago, Zanzibar, Dar-es- 
Salaam, Saya de Malha bank, Madras, Raiatea, Fiji, Tahiti, Rotuma, Samoa, Misol, 
New Guinea, Solomon Islands, Amboyna and Woosung. 

Richardson studied three specimens, two obtained by Sir James Ross during the 
voyage of H.M.S. " Erebus " and " Terror " (one of which was dissected) and another 
sent to the Royal College of Surgeons from Timor, where it had been found in a 
holothurian. The locality of the first two specimens is unknown, though Richardson 
believed it might be Tasmania. However, this species has not subsequently been 
reported further south than the latitude of New Caledonia. Only one of the three 
specimens is still in existence and this is in the collection of the British Museum 
(Natural History), no. 1952. 10. 30. 3. The type locality is generally stated as 
Tasmania but should probably be given as Timor. 

Specific characters. Maximum length c. 200 mm. ; length of head one-sixth to 
about one-seventh of total length ; maximum depth of head one-half to three-fifths 
and maximum width two-fifths to one-half of length of head ; specimens less than 
100 mm. long have relatively shorter and more slender heads than have larger specimens 
horizontal diameter of eye greater than length of snout, equal to interorbital width ; 
maxilla extends behind orbit to a distance equal to half horizontal diameter of eye ; 
one or two anterior teeth in upper jaw enlarged, the rest extremely small ; teeth of 
outermost series in lower jaw larger and stouter than those of inner series, bulging 
out a little beyond the edge of the jaw bone ; central row of usually three or four 
large, conical or slightly curved teeth on vomer, flanked by smaller ones ; pectoral 
fins half length of head ; anus anterior to roots of pectorals ; translucent in life 
with bluish or reddish shades anteriorly, dark cross-bars when adult (Jordan & 
Seale, 1905) ; silver spot between hind part of eye and maxilla in most specimens 
and a series of silver patches on flanks above lateral line. Oxybeles lumbricoides 
Bleeker is probably a tenuis larva of this species. 

Measurements and proportions of the type and of representative adults are given 
in Table VI, and a summary of the principal measurements of the adults studied in 
Table VII. 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 275 




E 
E 




CM 











'u 

a 

> 


'6 

a 

2 


ax 


CO 
4-> 


•rH 






X! 

c3 


O 


ef 


Ph 


cd 


>i 


(D 


+J 


O 


0) 


O 


■a 

H 


| 






<? 


T5 




1— 1 


HT 







4-» 


T3 


a 


Sh 





cd 


u 


^ 


SH 





a 


s 


0) 

a 











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§ 


0) 





CO 


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CO 




=s 




$, 




<S 




*v 




e 










276 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

Table VI. — Carapus homei {Richardson). Measurements and 
Proportions of the Type and of Representative Adults. 



Collection . 




British Museum 




Number 


1952. 10.30.3 


1875.3-3.8 


1908.3.23.36 


Locality- 


? 


Samoa 


Saya de Malha 




mm. 


mm. 


mm. 


Total length . 


109* 


. 142 


• 139 


Length of head 


16 (15% TL) 


20 (14% TL) 


22 (16% TL) 


Maximum depth of head 


9 (56% HL) 


10 (50% HL) 


12 (55% HL) 


Maximum width of head 


7 (44% HL) 


7 (35% HL) 


10 (45% HL) 


Length of snout 


3 (19% HL) 


4 (20% HL) 


4 (18% HL) 


Horizontal diameter of eye 


3-5(22%HL) 


5 (25% HL) 


5 (23% HL) 


Vertical diameter of eye 


3 (19% HL) 


4 -5(22%HL) 


4-5 (20% HL) 


Interorbital width 


2- 5 (i6%HL) 


4 (20% HL) 


4 (18% HL) 


Length of maxilla 


8-5( 53 %HL) 


11 (55% HL) 


n-5(52%HL) 


Length of pectoral fin 


8 (50% HL) 


13 (65% HL) 


10 (45% HL) 


Maximum depth of body 


9 (56% HL) 


13 (65% HL) 


15 (68% HL) 


Preanal length 


14 (13% TL) 


17 (12% TL) 


20 (14% TL) 



* Type specimen. 
TL = total length ; HL = length of head. 



Table VII. — Carapus homei (Richardson) . Summary of 
Measurements and Proportions of Adult Fish. 







R. 


M ± a M . 


Variate. 


N. 


mm. 


mm. 


Total length 


27 


100-190 


i38-6±5'09 


Length of head 


27 


15-30 


20-04^0-76 


Depth of head . 


27 


8-16 


IO-24jO-6l 


Width of head . 


27 


5-14 


8-i±o-6o 


Length of pectoral fin 


26 


7-15 


10-44^0-78 


Preanal length . 


26 


13-27 


I 7 , 7±o-8o 






% 


% 


Length of head (% TL) 


27 


12-8-17-0 


14-64^0-19 


Depth of head (% HL) 


27 


50 • 0-60 • 


54-3±o-i8 


Width of head (% HL) 


27 


32-0-50-0 


4 i-5±o-93 


Length of pectoral (% HL 


26 


40-0-65-0 


50-64^0-89 


Preanal length (% TL) 


26 


10-5-11-4 


I2-6±0-2I 



N = number of specimens ; R = range of variate ; 
mean o M ) ; TL = total length ; HL = length of head. 



M = mean of variate (i standard error of 



C. homei has a wide distribution and more extensive collections might show that 
it should be broken up into subspecies ; this, however, is not possible with the material 
at present available. It has been reported from Ghardaqa in the Red Sea, various 
Indian Ocean localities, Indonesian waters, the Philippines, Celebes, Torres Straits, 
off the coast of Queensland, Hawaii, Woosung, Palao and many localities in the 
South Pacific. It has been found mainly in species of Holothuria, Actinopyga and 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 277 

Stichopus, but also occasionally in other echinoderms (Culcita, Nardoa) and lamel- 
libranchs (Cardium, Pinctada) and once in an ascidian (Sty eld). 

Several nominal species have been described from within the range of C. hornet, 
which have differed from it solely in point of size and of number of vomerine teeth, 
neither of which, alone or together, can be regarded as good diagnostic characters. 
Two of these forms, Fierasjer brandesii Bleeker and Fierasfer sluiteri Weber, are now 
generally accepted as synonymous with C. homei, while two others, Fierasfer microdon 
Gilbert from Hawaii and Fierasfer mourlani Petit from Madagascar, are not distin- 
guishable in description from the smaller specimens of C. homei. None of these four 
forms has been recorded by other than the original authors, but two others, Fierasfer 
neglectus Peters from Mozambique and Fierasfer affinis Gunther of unknown origin, 
have been recorded upon a number of subsequent occasions and therefore merit 
more detailed consideration. 

F. neglectus was well described, though not figured, and there are in the British 
Museum collection a number of fierasfers from Indian Ocean localities that are 
clearly referable to this species. However, this group bears an equally close resemblance 
to the C. homei specimens from Indo- Australian and Pacific waters, especially in 
the dentition and the general presence of a suborbital silver patch. Specimens of 
similar size from the two areas have almost identical proportions. The two groups 
show small differences but comparison shows these differences to be too small 
for it to be impossible for them to have been drawn from the same " population ". 
F. neglectus may ultimately be found to be subspecifically distinct, but with the 
material at present available it is not possible clearly to diagnose to which group, 
Indie or Pacific, any given specimen belongs. 

F. affinis, of which the type only is known, has been regarded by most American 
authors as a fierasfer from American waters. This, however, is erroneous. It is 174 
mm. in length, far larger than any fierasfer yet recorded from the coastal waters of 
Central America ; the dentition is wholly that of C. homei ; there is a suborbital 
silver spot ; and its dimensions fit closely with those of the available sample of C. 
homei, especially those of the Pacific group. Comparison of the main characters of 
this specimen with those of the available sample of C. homei give values of d/cr which 
are well below 3-0, the minimum value usually taken as suggesting specific difference 
(Simpson & Roe, 1947). 

Carapus pindae Smith, 1955 
Carapus pindae Smith (J. L. B.), 1955, Ann. Mag. nat. Hist. (12) 8 : 412. 

The type, the only specimen known, was found inside an unidentified holothurian 
taken at Pinda (14 10' S., 40 40' E.) and is lodged in the Department of Ichthyology, 
Rhodes University, Grahamstown, South Africa. It has not been available for study 
and the following account is derived entirely from the original description. 

Specific characters. Total length 75 mm.; length of head one-seventh of 
total length ; width of head about one-third, depth of head about one-half, of length 
of head ; horizontal diameter of eye twice as great as interorbital width, one-half 
as great again as length of snout ; maxilla extending behind orbit for distance less 

zool. 4, 6. 19 



278 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

than one-half horizontal diameter of eye ; teeth in single series in upper jaw, the 
first pair enlarged ; teeth in bands in lower jaw, none enlarged ; palatine teeth 
uniserial posteriorly, polyserial anteriorly ; two large and several small teeth on 
vomer ; pectoral fins slightly less than one-half length of head ; anus anterior to 
roots of pectoral fins ; translucent in life with pink sheen, abdomen silvery-bronze, 
row of silvery-bronze spots on sides of abdomen, faint dusky cross-bars on back. 

The measurements of this specimen are not stated, but a series of proportions are 
given in the original description. 

This species is closest to C. hornet, but differs from it and all other fierasfers so 
far described in the partially uniserial nature of the dentition. Its small size suggests 
that C. pindae may not be fully mature, in which case the peculiarities of the dentition 
might be only transient. Should this indeed be the case, there seems little to justify 
separation of this species from C. homei as at present comprehended. 

Carapus kagoshimanus (Steindachner & Doderlein), 1877 

Fierasfer kagoshimanus Steindachner & Doderlein, 1877, Denkschr. Akad Wiss. Wien. 53 : 1. 

Jordan & Snyder, 1901, Annot. zool. jap. 3 : 118. 
Carapus kagoshimanus, Jordan & Fowler, 1903, Proc. U.S. nat. Mus. 25 : 743. Franz, 1910, 

Abhandl. Bayer Akad. Wiss., Suppl. 4:31. 

The type was obtained in the harbour at Kagoshima, Japan and its present 
whereabouts are unknown. No specimens have been available for study and the 
following account is based upon those of the authorities cited above. 

Specific characters. Length of type no mm.: head one-seventh of total 
length ; maximum depth of head three-fifths, maximum width two-fifths, of length 
of head ; horizontal diameter of eye equal to length of snout and to interorbital 
width ; teeth of jaws, palatines and vomer small ; anus beneath base of pectoral 
fins ; pectorals less than half length of head ; colour stated as uniform pink with 
dark spots. 

This species has been reported only from Kagoshima and Sagami . 

Carapus owasianus Matsubara, 1953 
Carapus owasianus Matsubara, 1953, J a P- J ■ Ichthyol. 3 : 29. 

The type, the only specimen known, was taken free-swimming off the coast near 
Owasi, Japan, and placed in Matsubara's Fish Collection, no. 18871. It has not been 
available for study and the following account is compiled from the original description. 

Specific characters. Total length about 200 mm. (determined from figure, not 
stated by author) ; head over one-eighth of total length ; horizontal diameter of 
eye a little greater than length of snout, nearly twice as great as interorbital width ; 
maxilla extending behind orbit for a distance approximately equal to half horizontal 
diameter of eye ; enlarged teeth at front of both jaws ; vomerine teeth small ; anus 
slightly posterior to roots of pectorals ; pectoral fins half length of head ; colour in 
life unknown. 

The measurements of this specimen have not been recorded, but its detailed 
proportions are listed in the original publication. 



SYSTEMATIC REVISION OF THE FAMILY TELEOST CARAPIDAE 279 

Carapus houlti (Ogilby), 1922 

Fierasfer houlti Ogilby, 1922, Mem. Queensland Mus. 7 : 301. 
Carapus houlti, McCulloch, 1929, Mem. Austral. Mus. 5 : 354. 

The two cotypes, the only specimens known, were taken together off Double 
Island Point, southern Queensland, in 36 fathoms (70 metres) enclosed in the remains 
of an unidentified holothurian. They have not been available for study and the 
following account is compiled from the original description. 

Specific characters. The two specimens were 283 mm. and 236 mm. long ; 
head one-seventh to one-eighth of total length ; width of head one-half its length ; 
maximum depth of body considerably greater than maximum depth of head ; teeth 
in jaws small and conical, uniform in size ; row of four large teeth on vomer ; anus 
beneath roots of pectorals ; pectoral fins a little less than one-third length of head ; 
colour greyish-brown with darker dots. 

Carapus boraborensis (Kaup), 1856 

Fierasfer boraborensis Kaup, 1856, Arch. Naturges. 22 : 97. Kaup, 1856, Catalogue of Apodal 

Fishes : 160. 
Jordanicus boraborensis, Jordan & Seale, 1905, Bull. Bur. Fish. 25 : 435. Fowler, 1928, Mem. 

Bishop Mus. 10 : 446. Fowler, 1938, Mon. Acad. nat. Sci., Philadelphia 2 : 260. 

The original description is based upon several specimens obtained by Lesson and 
Carnot at Borabora in the Samoan Archipelago and placed in the Paris Museum. 
They seem to be no longer in existence and no examples have been available for 
study. 

Specific characters. Kaup's two accounts are brief, unillustrated and in part 
contradictory. Iu the first the length is given as 330 mm., with the head one-eleventh 
of the total length and the pectoral fins one-third to one-quarter the length of the head; 
no other characters are mentioned. The second account records also the vomerine 
teeth and preanal length. In this description the largest specimen is said to be 23 in., 
or about 580 mm., in length ; the head is once again given as one-tenth to one- 
eleventh of the total length, but the pectoral fins are said to be one-sixth the length 
of the head. This fish has a cluster of thick teeth on the vomer and the anus is 
situated nearly 30 mm. anterior to the roots of the pectoral fins. 

This may not be a single species, nor even a fierasfer at all. It has been included 
in lists of the Samoan fish fauna (Jordan & Seale, 1905 ; Fowler, 1928 & 1938), 
but the citations are from Kaup and the species has not been seen since. There is 
no evidence to justify the inclusion of this rather doubtful form in Jordanicus. 

Carapus parvipinnis (Kaup), 1856 

(Text-fig. 12) 

Fierasfer parvipinnis Kaup, 1856, Arch. Naturges. 22 : 97. Kaup, 1856, Catalogue of Apodal 
Fishes : 160. Gunther, 1862, Catalogue of Fishes, 4 : 381. Fowler, 1900, Proc. Acad. nat. 
Sci., Philadelphia, 1900 : 523. Kendall & Goldsborough, 191 1, Mem. Mus. comp. Zool. 
Harvard, 26 : 330. Weber, 1913, Siboga Expeditie, 32 : 96. Tortonese, 1939, Boll. Mus. Zool. 
Anat. comp. Torino, 47 : 379. 



280 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 










v 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 281 

Jordanicus parvipinnis, Jordan & Seale, 1905, Bull. Bur. Fish. 25 : 435. Fowler, 1925. Proc. 

Acad. nat. Sci., Philadelphia, 77 : 264. Fowler, 1928, Mem. Bishop Mus. 10 : 447. Fowler, 

1938, Mon. Acad. nat. Sci., Philadelphia, 2 : 260. 
Carapus parvipinnis, Herre, 1936, Publ. Field Mus. zool. Ser. 21 : 416. Abe, 1939, Palao 

trop. biol. Stud. 1 : 575. Herre & Herald, 1950, Philipp. J. Sci. 79 : 337. Smith (J. L. B.), 

1955, Ann. mag. nat. Hist. (12) 8 : 412. 

The study material includes 12 adults in the collection of the British Museum 
(Natural History) and one in that of the Universitetets Zoologiske Museum, Copen- 
hagen. The localities represented are Tahiti, Ponape, Samoa, Rotuma, Raiatea, 
Banda and Mauritius. The type was obtained by Quoy and Gaimard at Carteret 
Harbour and probably placed in the Paris Museum. It seems to be no longer in 
existence. 

Specific characters. Maximum length about 300 mm.; length of head one- 
ninth to less than one-tenth of total length ; maximum depth of head about two- 
fifths and maximum width about one-half of length of head ; eye well below dorsal 
profile of head ; its horizontal diameter half as long again as length of snout, slightly 
less than interorbital width ; maxilla very stout, extending behind orbit for a 
distance equal to horizontal diameter of eye ; jaw teeth small, nearly uniform, a 
few at front of lower jaw smaller than rest ; three or four large, stout, conical teeth 
on vomer, surrounded by smaller teeth ; pectoral fins one-third to one-quarter 
length of head ; anus anterior to roots of pectorals. Of four specimens obtained 
alive at Moorea Island, Tahiti, two were a warm red-brown colour, the other two 
pale and translucent (Herre, 1 936ft). One reddish fish was certainly adult ; the pale 
fish may have been juveniles. 

Measurements and proportions of representative adults are given in Table VIII, 
and a summary for the adults examined in Table IX. 

Table VIII. — Carapus parvipinnis (Kaup). Measurements and 
Proportions of Representative Adults. 



Collection .... 




British Museum. 


Number .... 


1929.8.4. 1. 


. 


1874. 11. 19. 31. 


Locality .... 


mm. 




Tahiti. 


mm. 


Total length 


195 






260 


Length of head . 


21 


(n% TL) 






28 (11% TL) 


Maximum depth of head . 


13 


(62% HL) 






16 (57% HL) 


Maximum width of head . 


13 


(62% HL) 






12 (43% HL) 


Length of snout 


3 


(14% HL) 






4-5 (16% HL) 


Horizontal diameter of eye 


4 


(19% HL) 






5 (18% HL) 


Vertical diameter of eye . 


3'5 


(17% HL) 






4.5 (16% HL) 


Interorbital width . 


5 


(24% HL) 






7.5 (27% HL) 


Length of maxilla . 


10 


(48% HL) 






12.5 (45% HL) 


Length of pectoral fin . 


6 


(29% HL) 






8 (29% HL) 


Maximum depth of body 


14-5 


(69% HL) 






16 (57% HL) 


Preanal length . 


21 


(n% TL) 






27 (10% TL) 


TL = total lei 


lgth; HL 


= length of head 







282 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

Table IX. — Carapus parvipinnis (Kaup). Summary of 
Measurements and Proportions of Adults. 



Variate. 

Total length . 
Length of head 
Depth of head 
Width of head 
Pectoral length 
Preanal length 



Head length (% TL) 
Head depth (% HL) 
Head width (% HL) 
Pectoral length (% HL) 
Preanal length 

N = number of specimens ; R = r« 
HL = length of head. 

This species has been assigned (Jordan & Seale, 1905) to Jordanicus on account 
of the relatively great width of the head, even though it possesses neither of the 
other characters — adnate maxillae and absence of a lower lip — by which this genus 
was originally separated from Carapus. Instead, C. parvipinnis closely resembles 
C. acus in the form of its vertebrae, the shape of the lower jaw and the nature of the 
dentition. The most striking difference between this species and the other Carapus 
spp. is the relatively small size of the head which, when combined with the cylindrical 
body form, does cause it to approach Encheliophis {Jordanicus) gracilis in general 
appearance. However, this resemblance is purely superficial and provides no 
grounds for placing C. parvipinnis in any genus but Carapus. 

C. parvipinnis occurs in the tropical Pacific, especially at Tahiti, Samoa and the 
Philippine and Solomon Islands, as an inquiline of holothurians. One specimen 
in the collection of the British Museum was obtained in Mauritius. It has relatively 
longer pectorals than have the Pacific Ocean specimens, but this may be merely an 
individual peculiarity. 





R. 


M. 




mm. 


mm. 


10 


167-295 


224-9 


10 


18-33 


24-3 


10 


10-21 


14-8 


10 


8-21 


12-6 


10 


4-9 


6-9 


8 


18-33 


25-0 


10 


% 
9-55-11-67 


% 
io-8 


10 


55-56-65-38 


60-7 


10 


42 '10-63 '64 


5i-8 


10 


I9-05-37-50 


28-3 


8 


9-05-11-19 


io-6 


e; M = 


mean value of variate ; 


TL = total length 



Carapus caninus (Giinther), 1862 
(Text-fig. 13) 

Fierasfer caninus Giinther, 1862, Catalogue of Fishes, 4 : 381. Sauvage, 1891, Histoire Naturelle 

des Poissons : 476. 
Jordanicus caninus, Fowler, 1927, Bull. Bishop Mus., Honolulu, 38 : 30. Fowler, 1928, Mem. 

Bishop Mus. 10 : 447. Fowler, 1938, Mon. Acad. nat. Sci. Philadelphia, 2 : 260. 
Jordanicus fowleri, Smith (J. L. B.) 1955, Ann. Mag. nat. Hist. (12) 8 : 402. 
Carapus mayottae, Smith (J. L. B.) 1955, Ann. Mag. nat. Hist. (12) 8 : 415. 

The following account is based upon a re-examination of the type, no. 1952 . 10 . 30 . 2 
in the collection of the British Museum (Natural History) ; locality unknown. 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 283 

Specific characters. Total length of type is 83 mm.; body slender and strongly 
compressed, but not deeper than head ; head one-seventh of total length ; maximum 
depth of head one-half and maximum width one-third of length of head ; horizontal 
diameter of eye equal to length of snout, half as great again as interorbital width ; 
maxilla extends behind orbit for a distance equal to half horizontal diameter of eye ; 
outer row of teeth of lower jaw tall, curved, well-separated from each other, those 
at the front the largest ; inner teeth of lower jaw small, conical, close-set ; teeth of 




Fig. 13. — Carapus caninus (Giinther). 
vomerine teeth. 



2 mm 

The type and its 



upper jaw small, conical, pair at front greatly enlarged ; vomerine teeth in a single 
row as large as outermost teeth of lower jaw ; pectoral fins one-third length of head ; 
anus below roots of pectorals ; colour in life unknown. 
Measurement of this specimen are given in Table X. 

Table X. — Carapus caninus (Giinther). Measurements and 
Proportions of the type. 



Museum .... 


. British Museum. 


Number .... 


1952. 10.30.2. 




mm. 


Total length . 


88 


Length of head 


13 (15% TL) 


Maximum depth of head 


7 (54% HL) 


Maximum width of head 


4 (3i% HL) 


Length of snout 


3 (23% HL) 


Horizontal diameter of eye 


3 (23% HL) 


Vertical diameter of eye 


2.5 (19% HL) 


Interorbital width . 


2 (15% HL) 


Length of maxilla 


7-5 (57% HL) 


Length of pectoral fin 


5 (39% HL) 


Maximum depth of body . 


6 (46% HL) 


Preanal length . 


13 (15% TL) 


TL = total length ; HL = 


length of head. 



This species has also been recorded from Mayotte (Sauvage, 1891), host, if any, 
not stated ; and from Christmas Island (Fowler, 1927) when two specimens were 
taken in a pearl-oyster. 



284 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

Smith (1955), believing that the type is no longer in existence, has suggested that 
the trivial name caninus should be abandoned and has proposed instead the names 
Carapus mayottae for the specimen from Mayotte and Campus fowleri for those 
from Christmas Island. However, Carapus caninus is unquestionably valid and, 
though it has not been possible to examine the other specimens attributed to this 
species, the published descriptions and figures are not sufficiently at variance with 
Giinther's type to warrant specific separation. Fowler's inclusion of C. caninus in 
Jordanicus is also incorrect. 



Subgenus ONUXODON 

Subgeneric characters. Body strongly compressed, its greatest depth equal 
to length of head ; interorbital strongly domed, its width less than horizontal 
diameter of eye ; trunk vertebrae 19-20 ; swim-bladder partly calcified. 

Key to Species 
Pectoral fins less than one-quarter length of head . . C. (Onuxodon) parvibrachium. 

Pectoral fins more than one-half length of head . . C. (Onuxodon) margaritiferae. 



Carapus (Onuxodon) parvibrachium Fowler, 1927 
(Text-fig. 14) 

Carapus parvibrachium Fowler, 1927, Bull. Bishop Mus. 38 : 30. Fowler, 1928, Mem. Bishop 

Mus. 10 : 445. 
Onuxodon parvibrachium, Smith (J. L. B.), 1955, Ann. Mag. nat. Hist. (12) 8 : 406. 

The following account is based upon a single specimen obtained by Dr. T. Mortensen 
at Banda, Indonesia, and placed in the collection of the Universitetets Zoologiske 
Museum, Copenhagen. The holotype and one paratype were taken in Suva Bay, 
Fiji, in an unidentified clam-shell and placed in the Bernice P. Bishop Museum, 
Honolulu, Hawaii. 

Specific characters. Total length 67 mm. (type and paratype were 81 mm. and 
71 mm. respectively) ; length of head rather less than one-seventh of total length ; 
maximum depth of head a little less than three-quarters and maximum width two- 
fifths of length of head ; maximum depth of body equal to length of head ; hori- 
zontal diameter of eye slightly greater than length of snout, two and a half times as 
great as interorbital width ; maxilla extending behind orbit for a distance greater 
than horizontal diameter of eye ; anterior pair of teeth in each jaw very large, the 
rest of the dentition minute ; pectoral fins a little more than one-fifth length of head ; 
translucent in life with pink sheen and dark markings on snout, at bases of dorsal 
and anal fins and on caudal region (Smith, 1955). 

Measurements and proportions of this specimen are given in Table XI. 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 285 

Table XI. — Carapus (Onuxodon) parvibrachium Fowler. Measurements and 
Proportions of an Adult from Banda in the Collection of the Universitetets Zoologiske 
Museum, Copenhagen. 



Total length . 


67 




Length of head 


9 


(13% TL) 


Maximum depth of head 


7 


(78% HL) 


Maximum width of head 


3" 


5 (39% HL) 


Length of snout 


2 


(22% HL) 


Horizontal diameter of eye 


2- 


5 (28% HL) 


Vertical diameter of eye 


2 


(22% HL) 


Interorbital width . 


I 


(11% HL) 


Length of maxilla . 


6 


(67% HL) 


Length of pectoral fin 


2 


(22% HL) 


Maximum depth of body . 


9 


(100% HL) 


Preanal length . 


11 


(16% TL) 



TL = total length ; HL = length of head 




Fig. 14. 



10 mm 

-Carapus {Onuxodon) parvibrachium Fowler. A specimen from Bandu, Indonesia. 



Carapus (Onuxodon) margaritiferae (Rendahl), 1921 
(Text-fig. 15) 

Fierasfer homei Weber, 1913, Siboga Expeditie, 32 : 95. (non Richardson, 1844, Ichthyology of 
the Voyage of H. M.S. " Erebus " and " Terror " : 73.) 

Fierasfer margaritiferae Rendahl, 192 1, K. Svenska Vetensk. Akad. Handl. 61 (9) : 5. 

Carapus margaritiferae, de Beaufort & Chapman, 1951, Fishes of the Indo- Australian Archi- 
pelago, 9 : 449. Smith (J. L. B.), Ann. Mag. nat. Hist. (12) 8 : 400. 

? Fierasfer homei (part), Abe, 1939, Palao trop. biol. Stud. 1 : 574. 



286 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

The study material includes n adults in the collection of the Universitetets 
Zoologiske Museum, Copenhagen. Localities represented are Banda and Cape 
Jaubert. The type was taken from a pearl oyster dredged off Cape Jaubert, north- 
west Australia, and was placed in the Swedish Museum. 

Specific characters. Greatest recorded length 92 mm.: length of head one- 
sixth to one-seventh of total length ; maximum depth of head three-fifths and 
maximum width one- third of length of head ; horizontal diameter of eye less than 
length of snout, twice as great as interorbital width ; mouth nearly horizontal ; 
maxilla extends behind orbit for a distance greater than horizontal diameter of eye ; 
oval patch of many small teeth on vomer ; pectoral fins one-half to three-fifths of 
length of head ; translucent in life with pink sheen, caudal dusky (Smith, 1955). 




25mm 

Fig. 15. — Carapus (Onuxodon) margaritiferae (Rendahl). A representative specimen 

from Cape Jaubert, Australia. 



Measurements of representative specimens are given in Table XII, and a summary 
of measurements and proportions of the adult fish studied in Table XIII. 

Table XII. — Carapus (Onuxodon) margaritiferae (Rendahl). Measurements and 
Proportions of Representative Adults in the Collection of the Universitetets 
Zoologiske Museum, Copenhagen. 



Locality .... 


Banda. 


C. Jaubert. 




mm. 


mm. 


Total length 


68 


82 


Length of head . 


9 (13% TL) 


12 (15% TL) 


Maximum depth of head . 


5 (56% HL) 


8 (67% HL) 


Maximum width of head . 


3-5 (39% HL) 


5 (42% HL) 


Length of snout 


2 (22% HL) 


3 (25% HL) 


Horizontal diameter of eye 


1-3 (14% HL) 


1-5 (21% HL) 


Vertical diameter of eye . 


1 (11% HL) 


2 (17% HL) 


Interorbital width . 


0.7 (8%HL) 


i. 5 (i2%HL) 


Length of maxilla . 


5 (56% HL) 


7 ( 5 8% HL) 


Length of pectoral fin 


6 (67% HL) 


7 (58% HL) 


Maximum depth of body 


7 (78% HL) 


9.5 (79% HL) 


Preanal length . 


9-5 (14% TL) 


12 (15% TL) 


TL = total lei 


igth ; HL = length of head. 





SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 287 

Table XIII. — Carapus (Onuxodon), margaritiferae (Rendahl). 
Summary of Measurements and Proportions of Adults examined. 







R. 




M. 


Variate. 


N. 


mm. 




mm. 


Total length . 


10 


53-82 




68-8 


Length of head 


10 


5-12 




9*3 


Depth of head 


10 


3-8 




5'7 


Width of head 


9 


1-5-5 




3'3 


Length of pectoral fin 


9 


2-7 




5*6 


Preanal length 


9 


8-12 




9.7 


Length of head (% TL) 


10 


% 
8-93-15 


•38 


% 
13*23 


Depth of head (% HL) 


10 


50*00-71 


"43 


61-24 


Width of head (% HL) 


9 


30-00-57 


•14 


38-58 


Pectoral length (% HL) 


9 


40*00-71 


•43 


57*47 


Preanal length (% TL) 


9 


13.11-14-47 


13-59 


N = number of specimen ; R = rar 


lge of variate ; M = 


= mean value of variate ; 


TL = total length 


HL = length of head. 











C. margaritiferae has been found mainly in the Indo- Australian region, where it 
has been recorded from Cape Jaubert, Flores, Saleyer and Banda in the lamellibranchs 
Avicula and Pteria, in pearl oysters, and once in a holothurian. Five fierasfers 
taken at Palao (Abe, 1939) in Pinctada maxima may also belong to this species, 
while Smith (1955) has recorded it from clams at Durban. 



Carapus reedi Smith, 1955 

Carapus reedi Smith (J. L. B.), 1955, Ann. Mag. nat. Hist. (12) 8, p. 410. 

The type, the only specimen known, was found in a clam shell taken in 2 fathoms 
at Durban and is lodged in the Department of Ichthyology, Rhodes University, 
Grahamstown, South Africa. It has not been available for study and the following 
account is based entirely upon the original description. 

Specific characters. Total length about 70 mm.; head about one-fourteenth 
of total length ; maximum width of head about one-quarter, maximum depth 
nearly as great as length of head ; diameter of eye equal to length of snout, twice as 
great as interorbital width ; maxilla extends only as far as posterior edge of orbit ; 
jaw and palatine teeth small, uniserial ; anterior pair of teeth in each jaw enlarged ; 
vomerine teeth few, small ; pectoral fins two-thirds length of head ; anus posterior 
to base of pectorals ; translucent in life with pink sheen, few black spots on top of 
head. 

Measurements of this specimen have not been published, but detailed proportions 
are given in the original description. 

This specimen is clearly not a vexillifer larva, as are so many of the apparently 
aberrant fierasfers that have been described, but its small size, short maxilla, uniserial 
dentition and relatively short head form a combination of characters which suggest 
that it has not yet attained full maturity. From the illustration it is undoubtedly 



288 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

a Carapus, while the proportions of the head suggest that it should be placed in the 
subgenus Onuxodon. If it is indeed a late tenuis larva or early juvenile, it is probably 
attributable to the preceding species, C. margaritiferae. 



Genus ECHIODON Thompson, 1837 

Genotype Echiodon Drummondii Thompson, 1837 

Echiodon Thompson, 1837, Proc. zool. Soc. Lond. 5 : 55. Type species Echiodon Drummondii 

Thompson, 1837. 
Ophidium (Echiodon), Thompson, 1841, Trans, zool. Soc. Lond. 2 : 207. 

Generic characters. Body elongated, cylindrical ; lateral processes of first 
and second vertebrae expanded, not fused ; processes of all other vertebrae small 
and not expanded ; trunk vertebrae number 27-28 ; lower jaw stout, nearly straight, 
narrowing at distal end, then expanding again at extreme tip ; one or two pairs of 
large, fang-like teeth at front of both jaws ; other jaw teeth small and close-set in 
narrow bands ; anterior fangs of lower jaw separated from other teeth by a large 
diastema ; maxilla extending beyond orbit in adult, clearly outlined by folds of 
skin ; anus posterior to roots of pectorals in adult ; upper edge of orbit impinging 
on dorsal profile of skull ; swim-bladder long and straight, not constricted ; branchio- 
stegals 7. 

The genus Echiodon was first created to contain the newly discovered species 
E. drummondi. Subsequently, its author reduced it to subgeneric rank under 
Ophidium and Kaup (1856^) united it with Fierasfer. Later authors have followed 
Kaup, but the differences between E. drummondi and most other fierasfers are so 
clear-cut that it is necessary to re-establish the genus, not only for the original 
species, but also for that originally described as Ophidium dentatum Cuvier (1817). 

Echiodon differs from both Carapus and Encheliophis in both vertebral and jaw 
structure, in the clearly posterior position of the anus in the adult and in the presence 
of a diastema at the front of both jaws. It is further distinguished from Encheliophis 
in the possession of polyserial dentition and from Carapus in the relative shortness 
of the head. 

Key to Species, Adults only 

Upper profile of head rounded vomer tapering posteriorly ; no gap between vomerine 

and palatine teeth .......... E. drummondi 

Upper profile of head straight ; vomer rounded posteriorly ; distinct gap between 

vomerine and palatine teeth . . . . . . . . . E. dentatus 



Echiodon drummondi Thompson, 1837 
(Text-fig. 16) 

Echiodon drummondii Thompson, 1837, Proc. Zool. Soc. Lond. 5 : 52. Yarrell, 1852, Proc. Zool. 

Soc. Lond. 20 : 14. Edward, 1863, Zoologist (1) 21 : 8495. Couch, 1864, History of the Fishes 

of the British Isles, 3 : 133. 
Ophidium (Echiodon) drummondii, Thompson, 1841, Trans, zool. Soc. Lond. 2 : 207. 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 289 

Fierasfer dentatus Kaup, 1856, Catalogue of Apodal Fish : 157. (non Ophidium dentatum 
Cuvier, 1817, Regne Animal, 2 : 239.) Giinther, 1862, Catalogue of Fishes, 4 : 381. Day, 
1880, The Fishes of Great Britain and Ireland, 1 : 228. Colett, 1882, Forh. VidenskSelsk. y 
Kristiania, 1882, (19) : 5. Sim. 1883, Scottish Nat., 7 : 55. Fries, Ekstrom & Sundevall, 
1893, History of Scandinavian Fishes. 2 : 260. Aflalo, 1904, British Salt-water Fish : 294. 
Ehrenbaum-Helgoland, 1909, Nord Plank. 10 : 217. Grieg, 191 1, Bergen Mus. Aarbok. 
1911 (6) : 17. Ehrenbaum-Helgoland, 1936, Naturgeschichte und Wirtschaftliche Bedeutung 
der Seefische Nordeuropas 115. 

The following account is based upon 132 adults in the collection of the Scottish 
Home Department's Laboratory at Torrey, Aberdeen, obtained mainly from the 
coastal waters of north and west Scotland and Northern Ireland, and three late 
vexillifers in the collection of the British Museum (Natural History), obtained at 
Banff. The type was found dead on a beach at Glenarm, Co. Antrim, Northern 
Ireland. It can no longer be traced. 




50 mm 



Fig. 16. — Echiodon drummondi Thompson. A representative specimen 
from Scottish waters. 



Specific characters. Greatest recorded length 300 mm.; length of head one- 
ninth to one-eleventh of total length ; maximum depth of head about one-half and 
maximum width about one-third of length of head ; upper profile of head convex ; 
horizontal diameter of eye slightly greater than length of snout, about twice as great 
as interorbital width ; maxilla extends well behind posterior edge of orbit ; anus 
3-5 mm. posterior to roots of pectoral fins ; pectorals about three-fifths length of 
head ; patch of teeth on vomer rounded anteriorly, tapering posteriorly and merging 
with bands of palatine teeth ; colour in life believed to be reddish, with silvery 
abdomen, operculum and iris and dark markings on top of head, along edges of 
median fins and on tip of tail. (Silvery regions and dark markings are usually visible 
in preserved specimens.) 

There is a vexillifer larva, resembling adult in form of lower jaw and in dentition. 
It is translucent in life, with reddish colour along dorsal and ventral surfaces, silvery 
spot near anus, dark-green pupil and silvery iris (Edward, private communication, 
cited by Couch, 1864). Eggs and other stages in the life history are unknown. 

Measurements and proportions of representative adults are given in Table XIV, 
and a summary for the specimens examined in Table XV. 



2 9 o SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

Table XIV. — Echiodon drummondi Thompson. Measurements 
and Proportions of Representative Adults. 







6i°oi , N., 


oo° 30' 


W. 




Locality. 




in 155 m. 




Adriatic. 




mm. 




mm. 




mm. 


Total length . 


215 




250 




200 


Length of head 


. 24 


(11% TL) 


• 25 


(10% TL) 


. 22-5 (11% TL) 


Maximum depth of head 


10 


(42% HL) 


12 


(48% HL) 


10 (44% HL) 


Maximum width of head 


8 


(33% HL) 


9 


(36% HL) 


7 (3i% HL) 


Length of snout 


4 


(15% HL) 


6 


(24% HL) 


4-5 (20% HL) 


Horizontal diameter of eye 


5 


(21% HL) 


6 


(24% HL) 


5 (22% HL) 


Vertical diameter of eye 


4 


(17% HL) 


4-5 


(18% HL) 


4 (18% HL) 


Interorbital width 


2-5 


(10% HL) 


4 


(16% HL) 


4 (18% HL) 


Length of maxilla 


10.5 


(44% HL) 


13 


(52% HL) 


11-5 (51% HL) 


Length of pectoral fin 


14 


(58% HL) 


• 15 


(60% HL) 


13 (58% HL) 


Maximum depth of body 


10 


(42% HL) 


12 


(48% HL) 


10 (44% HL) 


Preanal length 


26 


(12% TL) 


32 


(13% TL) 


24 (12% TL) 



TL = total length ; HL = length of head. 



Table XV. — Echiodon drummondi Thompson. Summary of 
Measurements and Proportions of Adults. 



N 



Variate. 


N. 


R. 

mm. 


M ± a M • 
mm. 


Total length 
Length of head 
Depth of head . 
Width of head . 


104 
114 
114 
108 


160-292 

15-30 

8-i 5 

6-13 


257'i±2-59 

25-4±o-26 

12-1^0-14 

9-7±o-i4 


Pectoral length 
Preanal length . 


114 
112 


10-20 
20-38 


i5-o±o-i9 
3o*7±o-34 


Length of head (% TL) 
Depth of head (% HL) 
Width of head (% HL) 
Pectoral length (% HL) 
Preanal length (% TL) 


104 

114 
108 
114 
102 


% 
9 -09-11-73 
40-00-56-00 
23-26-46-43 
41-27-46-43 
11 -11-14-80 


% 
10-31^0-05 
47-48^0-28 
3 8-i 4 ±o-36 
5 8.8 9 ±o- 5 6 
12-48^0-04 


number of specimens ; R = 
o M ) ; TL = total length ; HI 


range of variate ; 
, = length of head. 


M = mean of var 


iate (± standard e 



Echiodon drummondi is apparently not uncommon in the coastal waters round the 
north and west coasts of the British Isles and has been taken at a considerable 
number of localities at depths down to 100 fathoms (200 metres). It has also been 
reported from Scandinavian waters. A specimen from the Oceanographic Institute 
at Split is also assigned to E. drummondi, though it may ultimately prove to be 
subspecifically distinct. The measurements of this specimen are given in Table XIV. 

Cuvier described two fierasfers from the Mediterranean of which one, Ophidium 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 291 

imberbe, is that now termed Carapus acus. The other, Ophidium dentatum Cuvier, 
1817, was said to differ from the first and more common species only by the presence 
of " deux dents en crochets ", a character which barely constitutes a recognizable 
description. Kaup (18560) examined a number of specimens of 0. dentatum, at that 
time preserved in the Paris Museum, and concluded that Cuvier's Mediterranean 
and Thompson's Atlantic fierasfer were identical. Both have therefore been recorded 
by later authors as Fierasfer dentatus, the name used by Kaup. 

Though C. acus is fairly common in the Mediterranean, the other fierasfer is 
extremely rare and known from only a few localities. I have been able to examine 
only two, one from Monaco and another, very poorly preserved, from Sicily. The 
Atlantic fierasfer is also generally regarded as extremely rare and has even been 





1 1 

20 mm 

Fig. 17. — Heads of Echiodon spp. a, E. dentatus ; b, E. drummondi. 



mentioned by some authors as an exotic stray from the Mediterranean ; but during 
the past fifty years (excluding the war periods) the research vessels of the Aberdeen 
Laboratory have obtained well over 100 adult fish which accord well with the descrip- 
tion given by Thompson. 



292 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

It is almost certain that Cuvier's types, assuming they were indeed the specimens 
seen by Kaup, are no longer in existence ; certainly they are not in the Paris collec- 
tion. Thompson's specimen has also disappeared. It is therefore impossible to 
determine the status of these two forms by comparison of types, but from examina- 
tion of the specimens preserved at Aberdeen and Monaco conforming to the descrip- 
tion of E. drummondi and 0. dentatum respectively it is clear that they are in fact 
two different species. They closely resemble each other in the structure of the lower 
jaw and of the first three vertebrae, in the presence of fang-like teeth at the front of 
both jaws and in the posterior position of the anus. However, they differ greatly 
in their dimensions and to a certain extent in their proportions. Other differences 
are apparent in the shape of the head, the obliquity of the mouth and the form of 
the operculum (Text-fig. 17). A further difference lies in the shape of the vomer, as 
displayed by the teeth it bears. In the Atlantic form the vomer tapers posteriorly, 
and there is no gap between vomerine and palatine teeth ; in the Mediterranean 
species the vomer terminates abruptly and its teeth are separated from those of the 
palatines by a well-marked gap. 



Echiodon dentatus (Cuvier), 1817 
(Text-fig. 18) 

Ophidium dentatum Cuvier, 181 7, Regne Animal, 2 : 239. 

Fierasfer dentatus, Kaup, 1856, Verh. phys.-med. Ges. Wurzburg. 7 : 233. Kaup, 1856, Arch. 
Naturges, 22 : 93. Giglioli, 1880, Esposizione di Pesca : 97. Emery, 1880, Fauna u. Flora 
Neapel, 2 : 17. Carus, 1885, Prodromus Faunae Mediterranea, 2 : 580. Raffaele, 1888, Mitt, 
zool. Stat. Neapel, 8:1. Moreau, 1891, Histoive Naturelle des Poissons de la France, Supple- 
ment : 59. Belotti, 1891, Atti Soc. Hal. Sci. nat. Milano, 33 : 127. Moreau, 1892, Manuel 
d'Ichthyologie Francaise : 405. 

The following account is based upon one specimen in the collection of the Institut 
Oceanographique, Monaco ; locality, Monaco : and another in the collection of the 
Universitetets Zoologiske Museum, Copenhagen ; locality Sicily. 

Cuvier stated that his fish came from the Mediterranean. He did not designate a 
type and it is not certain that his specimens were preserved. The specimens examined 
by Kaup came, he believed, from Naples, but seem to be no longer in existence. 

Specific characters. Total length 170 mm.; length of head one-ninth of total 
length ; upper profile of head straight ; maximum depth of head two-fifths, and 
maximum width one-third, of length of head ; horizontal diameter of eye slightly 
less than length of snout, nearly twice as great as interorbital width ; anus 4 mm. 
posterior to roots of pectorals ; pectoral fins slightly less than one-half length of 
head ; maxilla extends behind eye for distance equal to horizontal diameter of eye ; 
vomer rounded posteriorly ; vomerine and palatine teeth separated by a distinct 
gap ; colour in life unknown. Eggs attributable to this species have been described 
(Raffaele, 1888). 

Measurements of this specimen are given in Table XVI. Echiodon dentatus has 
been recorded from Nice, Monaco, Naples, Sicily and the Adriatic. 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 



293 



ttl 



ZOOL. 4, 6. 



294 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

Table XVI. — Echiodon dentatus (Cuvier). Measurements and Proportions of a 
Specimen in the Collection of the Institut Oceanographique de Monaco. 



Total length 


170 




Length of head 


19 


(11% TL) 


Maximum depth of head 


8 


(42% HL) 


Maximum width of head 


6 


(32% HL) 


Length of snout 


3'5 


(18% HL) 


Horizontal diameter of eye 


3*2 


(17% HL) 


Vertical diameter of eye 


2-5 


(14% HL) 


Interorbital width . 


2 


(11% HL) 


Length of maxilla . 


10 


(53% HL) 


Length of pectoral fin 


9 


(47% HL) 


Maximum depth of body . 


9 


(47% HL) 


Preanal length . 


23 


(14% TL) 


TL = total length ; HL = 


length of head. 



SPECIES POSSIBLY ATTRIBUTABLE TO ECHIODON 

Carapus rendahli Whitley, 1941 

Fierasfer sp., Rendahl, 1925, Vidensk. Medd. Dansk. Nat. Foren. 81 : 13. 
Carapus rendahli Whitley, 1941, Austral. Zool. 10 : 40. 

The type, the only specimen known, was obtained at Port Jackson, New South 
Wales, and is now in the Australian Museum, no. 1 . 2, 411. It has not been available 
for study and the following account is compiled from Whitley (1941). 

Specific characters. Total length 93 mm.; length of head less than one-eighth 
of total length ; maximum depth of head a little less than one-half and maximum 
width one-quarter of length of head ; horizontal diameter of eye as great again as 
length of snout, three times interorbital width ; maxilla extends a short distance 
behind orbit ; anus about 3 mm. posterior to roots of pectorals ; pectoral fins 
one-quarter length of head ; colour in life unknown. A number of vexillifers from 
New South Wales waters were provisionally attributed to this species. 

The specimen is small and so may not be fully adult. The tip of the tail is broken 
off and a portion of the intestine protrudes through the anus. The general appearance 
as figured by Whitley is not that of a Carapus, but of a fish closely resembling 
Echiodon drummondi. The correspondence with Echiodon is shown also by the presence 
of fang-like teeth at the front of the jaws and by the posterior position of the anus, 
but cannot be proved without reference to the vertebral and jaw structure, neither 
of which have been described. It is not clear from the figure whether this specimen 
has a diastema in the lower jaw. Two further points suggesting that C. rendahli may 
in fact be an Echiodon are provided by the broken tail and intestinal prolapse. 
Damage to the tail is frequent in Echiodon and nearly 10% of the E. drummondi 
in the Aberdeen collection have broken or regenerating tails. On the other hand, 
Carapus spp. have seldom been found with broken tails, never with the tail stump 
showing signs of regeneration. Prolapse is also frequent in preserved Echiodon 
drummondi, unknown in Carapus spp. These two characters are not, of course, 
diagnostic, but are extremely suggestive. 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 295 

Carapus cinereus Smith, 1955 
Carapus cinereus Smith (J. L. B.), 1955. Ann. Mag. nat. Hist. (12) 8 : 409. 

The type, the only specimen known, was taken in a tide pool at Inhaca Island 
(26 S., 33 D.) and is preserved in the Department of Ichthyology, Rhodes University, 
Grahamstown, South Africa. It has not been available for study and the following: 
account is derived entirely from the original publication. 

Specific characters. Total length 215 mm.; length of head less than one-ninth 
of total length ; maximum width of head two-fifths, maximum depth about one-half 
of length of head ; diameter of eye less than length of snout, greater than interorbital 
width ; maxilla extends behind eye for distance equal to about one-half diameter of 
orbit ; teeth of jaws small, blunt, conical, arranged in bands which terminate well 
before symphysis ; large, curved teeth at front of both upper and lower jaws, from 
description apparently separated by a diastema from other teeth ; vomerine teeth 
conical, of several sizes ; palatine teeth conical, generally small ; pectoral fins 
about one-half length of head ; anus clearly posterior to base of pectorals ; translu- 
cent in life with pink sheen, abdomen silvery, top of head dusky. 

Apart from the total length the measurements of this specimen have not been 
published, but proportions are listed in the original paper. 

The extreme length, relatively short head and cylindrical body of this specimen 
are characters which, though shown to a certain extent by C. parvipinnis, are by 
no means typical of the genus Carapus. The nature of the lower jaw and dentition 
are sufficient to exclude this species from Encheliophis, and its general appearance 
and the presumed presence of a diastema between the large anterior teeth and small 
posterior teeth in both jaws suggest that in fact this species should be assigned to 
Echiodon. The posterior position of the anus, the shape of the head, flatness of the 
interorbital and impingement of the upper edge of the orbit upon the dorsal profile 
of the head are further characters in which this form differs from veritable Carapus 
spp. and resembles E. drummondi. 

Genus ENCHELIOPHIS Miiller, 1842 
Type species Encheliophis vermicularis Miiller, 1842 

Encheliophis Miiller, 1842, Ber. Verh. Preuss. Akad. 1842 : 205. Type species Encheliophis 

vermicularis Miiller, 1842. 
Jordanicus Gilbert, 1905, Bull. U.S. Fish. Comm. 23 : 655. Type species Fierasfer umbratilis 

Jordan & Evermann, 1902, Bull. U.S. Fish. Comm. 22 : 206. 
Encheliophiops Reid, 1940, Rep. Allan Hancock Pacific Exp. 9 : 47. Type species Enchelio- 

phiops hancocki Reid, 1940. 

Generic characters. Body elongated, cylindrical ; lateral processes of first 
and second vertebrae long, not expanded ; of third, fourth and fifth vertebrae long, 
expanded and fused into broad plates ; of sixth and subsequent vertebrae short, 
not expanded ; trunk vertebrae 30-31 ; lower jaw slender, curved, tapering to tip ; 
teeth in single rows on jaws and palatines even in adult ; no diastema ; no fleshy 
lip to lower jaw ; maxilla concealed beneath skin ; interorbital domed, raising upper 



296 SYSTEMATIC REVISION OF THE FAMILY TELEOST CARAPIDAE 

profile of skull considerably above orbit ; pectorals small or absent, but pectoral 
girdle always present ; anus beneath roots of pectoral in adult ; branchiostegals 
6 or 7. 

This genus was created to contain a finless fierasfer, Encheliophis vermicularis, 
from the Philippines. Subsequently other finless forms have been described from 
localities off the Pacific coast of Central America. One of these species, E. hancocki, 
introduced a new generic name Encheliophiops. This genus was separated from 
Encheliophis solely on the grounds that the only specimen known had the tip of the 
tail finless. This character, however, is shown by certain of the tenuis larvae of 
Carapus acus and by itself cannot be regarded as adequate for generic separation. 
Encheliophiops is accordingly regarded as synonymous with Encheliophis. 

Also included with Encheliophis in the present work are the species customarily 
assigned to the genus Jordanicus. This genus was created to contain the single 
species Fierasfer umbratilis (generally accepted as a synonym for Oxybeles gracilis) 
and was stated to differ from other genera in the uniserial dentition, concealed maxilla 
and absence of a lip in the upper jaw. Radiographs of the type and other specimens 
of Oxybeles gracilis have added the structure of the lower jaw and fusion between 
the 3rd, 4th and 5th vertebrae to the original characters. In all these features the 
species assigned to Jordanicus differ from all other fierasfers except species of Encheli- 
ophis. The only difference which it has been possible to detect between Encheliophis 
and Jordanicus is the absence in the former of pectoral fins. However, the pectoral 
fins of Jordanicus are relatively far smaller than those of Carapus spp. and obvious 
though the presence of pectoral fins is as a diagnostic character, it is considered that 
this difference, unsupported by other characters, is insufficient to warrant generic 
distinction. The two genera are accordingly united, Encheliophis Miiller, 1842, 
taking precedence over Jordanicus Gilbert, 1905. Jordanicus is retained, however, 
as a subgenus under Encheliophis. 



Subgenus ENCHELIOPHIS 

Subgeneric characters. Pectoral fins absent ; branchiostegals 6. 

Key to Species 

1 . Membranes of dorsal and anal fins continuous round tip of tail .... 2 
Membranes of dorsal and anal fins not continuous round tip of tail . . E. hancocki 

2. Body darkly pigmented . . . . . . . . . E. vermicularis 

Body not darkly pigmented . . . . . . . . E. jordani 

Encheliophis vermicularis Miiller, 1842 
(Text-fig. 19) 

Encheliophis vermicularis Miiller, 1842, Ber. Verh. preuss. Akad. Wiss. 1842 : 205. Miiller, 1843 

Abhandl. Akad. Wiss. Berlin, 1843 : 109. Kaup, 1856, Catalogue of Apodal Fish : 157. 

Giinther, 1862, Catalogue of Fishes, 4 : 381. Herre, 1936, Publ. Field Mus. zool. Ser. 21 : 417. 

Abe, 1939, Palao trop. biol. Stud. 1 : 574. Smith (J. L. B.), 1955, Ann. Mag. nat. Hist. (2) 

8 : 416. 
Enchelyophis vermicularis Miiller, 1843, Arch. Naturges. 9 : 329. 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 297 

The study material includes a single specimen in the collection of the Universitetets 
Zoologiske Museum, Copenhagen, from Sambelong, mouth of the Ganges ; and two 
in the British Museum (Natural History) obtained off Lizard Island, Great Barrier 
Reef, and at Tahiti. The type was obtained in the Philippines. It is uncertain 
whether it is still in existence. 

Specific characters. The following description is of the specimen obtained at 
Sambelong by the Galathea expedition and placed in the Universitetets Zoologiske 
Museum. 

Total length 79 mm. (greatest recorded for this species 175 mm.) ; head one- 
eleventh of total length ; maximum depth of head two-fifths and maximum width 
rather less than one-third of length of head ; horizontal diameter of eye slightly 




20 mm 

Fig. 19. — Encheliophis vermicularis Miiller. A specimen from 
Sambelong, India. 

less than length of snout, slightly more than interorbital width ; maxilla extends 
behind orbit for a distance equal to half horizontal diameter of eye ; preanal length 
equals length of head ; dark brown in life (Miiller, 1842). Life history unknown. 
Measurements of this and another specimen are given in Table XVII. 



Table XVII. — Encheliophis vermicularis Miiller. Measurements and Prof 


Collection . 


Danish 


British 




Museum. 


Museum. 


Number 


— 


1933.8. 12.47. 


Locality 


Sambelong. 


Lizard Is. 




mm. 


mm. 


Total length 


79 


134 


Length of head 


7 (9% TL) 


11. 5 (9% TL) 


Maximum depth of head 


3 (43% HL) 


5-5(48%HL) 


Maximum width of head 


2 (29% HL) 


4 (35% HL) 


Length of snout 


1.3 (19% HL) 


2 (17% HL) 


Horizontal diameter of eye 


1. 1 (16% HL) 


2 (17% HL) 


Vertical diameter of eye 


1 (14% HL) 


1.7 (17% HL) 


Length of maxilla 


3 (43% HL) 


6 (52% HL) 


Interorbital width 


1 (14% HL) 


2 (17% HL) 


Maximum depth of body 


3-5 (50% HL) 


6 (72% HL) 


Preanal length 


7 (9% TL) 


12 (9% TL) 



TL = total length ; HL = length of head. 

E. vermicularis has been recorded from the coasts of Somaliland and India, the 
Philippines, Palao, Sulu Sea and Tahiti. It was taken in holothurians in the Philip- 
pines and is said to feed upon the viscera of its host (Semper, 1861). 



298 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

Encheliophis jordani Heller & Snodgrass, 1903 
Encheliophis jordani Heller & Snodgrass, 1903, Proc. Acad. Sci. Washington, 5 : 220. 

The type, the only specimen known, was obtained at Tagus Cove, Albermarle 
Island, during the Hopkins Stanford Galapagos Expedition and deposited in Leland 
Stanford Junior University Museum, catalogue no. 6345. It has not been available 
for study and the following account is based upon the original description. 

Specific characters. Total length 114 mm.; length of head one-eleventh of 
total length ; maximum depth of head one-half its length, maximum width not 
stated ; horizontal diameter of eye less than snout length, slightly more than inter- 
orbital width ; maxilla extending short distance behind orbit ; small, rounded, 
patch of teeth on vomer ; anus below posterior border of operculum ; head and 
body dusky pink in life, abdomen silver, tail greyish-lavender, iris greenish-grey ; 
colours fading on preservation. Life history unknown. 



Encheliophis hancocki (Reid), 1940 

Encheliophiops hancocki Reid, 1940, Rep. Allan Hancock Pacific Exp. 9 : 47. Steinbeck & 
Ricketts, 1941, Sea of Cortez : 575. 

The type was taken among the coral Pocillipora sp. at Gorgona Island, Columbia, 
during the course of the Hancock Pacific Expedition, 1935, and placed in the United 
States National Museum, catalogue no. 101789. It has not been available for study. 

Specific characters. Total length of type 74*8 mm. ; length of head one- 
eleventh of total length ; maximum depth one-half and maximum width two-fifth 
of length of head ; horizontal diameter of eye equal to length of snout, slightly 
greater than interorbital width ; maxilla extends slightly behind orbit ; vomerine 
teeth not described ; anus about 2 mm. behind hind border of operculum ; median 
fins not continuous round tip of tail, which ends in a bare point : colour in life 
unknown. Life history unknown. 

A second specimen was obtained from the cloaca of Holothuria lubrica taken in 
the Gulf of California (Steinbeck & Ricketts, 1941). 

The status of the species of Encheliophis is doubtful and it is not entirely clear to 
which the various recorded specimens ought to be assigned. Certainly there is one 
fierasfer, Encheliophis vermicularis, which lacks paired fins, but though the majority 
of specimens noted by other authors have been ascribed to this species, they have 
not been figured, nor have their measurements been recorded. In consequence it 
has not been possible to build up a body of data giving some picture of the species 
as a whole, as has been the case with other members of the Carapidae. 

The species of Encheliophis do not, so far as is at present known, differ greatly 
in proportions or in dentition and separation is based mainly upon colour. This is 
not a good criterion, but is the best available. Certainly, the dark pigmentation of 
E. vermicularis, which persists on preservation, makes it easy to distinguish this 
species from the paler E. jordani. E. jordani and E. hancocki resemble each other 
very closely indeed, and may ultimately prove to be conspecific. 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 299 

Subgenus JORDANICUS 
Subgeneric characters. Pectoral fins present, small ; branchiostegals 7. 

Key to Species (Adults only) 

Vomerine teeth in single short median row . . . . . E. (Jordanicus) gracilis 

Vomerine teeth in short band of about four rows . . E. {Jordanicus) sagamianus 

Encheliophis (Jordanicus) gracilis (Bleeker), 1856 
(Text-fig. 20) 

Oxybeles gracilis Bleeker, 1856, Nat. Tijdschr. Ned.-Ind. 11 : 105. Doleschall, 1858, Nat. 

Tijdschr. Ned.-Ind. 15 : 163. 
Fierasfer gracilis, Giinther, 1862, Catalogue of Fishes, 4 : 381. Pietschmann, 1938, Bull. 

Bishop Mus. 156 : 51. Tortonese, 1939, Boll. Mus. zool. Anat. comp. Torino, 47 : 379. 
Fierasfer punctatus Fischer, 1885, Jahrb. Hamburg Wiss. Anst. 2 : 74. Barnard, 1927, Ann. S. 

Afr. Mus. 21 : 884. 
Fierasfer umbratilis Jordan & Evermann, 1902, Bull. U.S. Fish. Comm. 22 : 206. Jordan & 

Evermann, 1903, Bull. U.S. Fish. Comm. 23 : 505. 
Jordanicus umbratilis Gilbert, 1905, Bull. U.S. Fish. Comm. 23 : 655. 
Fierasfer frantii Popta, 19 12, Notes Ley den Mus. 23 : 185. 
Carapus gracilis, Fowler, 1925, Proc. Acad. nat. Sci. Philadelphia, 77 : 283. Fowler, 1926, Ann. 

Natal Mus. 5 : 402. de Beaufort & Chapman, 1951, Fishes of the Indo- Australian Archi- 
>, 9 : 449. 




25 mm 

Fig. 20. — Encheliophis {Jordanicus) gracilis (Bleeker). 
a, The type ; b, a more fully adult specimen. 

Fierasfer {Jordanicus) gracilis, Barnard, 1927, Ann. S. Afr. Mus. 21 : 884. 

Jordanicus gracilis, Fowler, 1928, Mem. Bishop Mus. 10 : 447. Fowler, 1931, Mem. Bishop Mus. 
11 : 364. Fowler, 1934, Mem. Bishop Mus. 11 : 447. Abe, 1939, Palao trop. biol. Stud. 
1 : 575. Schultz, 1943, Bull. U.S. nat. Mus. 180 : 287. Smith (J. L. B.), 1949, Sea Fishes 
of Southern Africa : 359. Smith (J. L. B.), 1955, Ann. Mag. nat. Hist. (12) 8 : 404. 

Carapus puntatus, Smith (J. L. B.), 1949, Sea Fishes of Southern Africa : 359. 

Jordanicus punctatus, Smith (J. L. B.), 1955, Ann. Mag. nat. Hist. (12) 8 : 405. 

Study material includes the type and nine other adults in the collection of the 
British Museum (Natural History) ; localities represented are Samoa, Pelew Islands, 



300 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

Tahiti. Varaii, Ovalau, Savaii and Wallis Island ; also one adult in the collection of 
the Universitetets Zoologiske Museum, Copenhagen, from Ghardaqa, Red Sea. 

The type was obtained at Banda Island, Indonesia, and is now in the collection of 
the British Museum (Natural History) under catalogue no. 1861.2.28.51. 

Specific characters. Greatest recorded length 236 mm.; length of head one- 
eighth to one-tenth or less of total length ; maximum depth of head about two-fifths 
of total length, maximum width slightly less ; horizontal diameter of eye equal to 
length of snout, slightly greater than interorbital width ; mouth almost horizontal ; 
jaw teeth well-separated from each other ; posterior part of lower jaw toothless in 
some specimens ; vomerine teeth one to four, in single median row, conical and of 
moderate size ; pectoral fins one-third length of head. 

The colour of the living animal has been described upon four occasions, but the 
accounts are conflicting. Jordan and Evermann (1902) described a fish taken at 
Hilo as pale olivaceous with greenish spots. The colour pattern is given with meticulous 
detail, not only for the body, but also for the first and second dorsal, anal, caudal, 
pectoral and pelvic fins. Obviously, these notes do not refer to a fierasfer at all. The 
next description (Fowler, 1925) is of a specimen from the Natal Coast, recorded as 
" ecru-drab with darker mottlings", while in a subsequent paper (Fowler, 1926) 
the same specimen was described as pale mauve with a blue tinge. Smith (1955) 
described a specimen from Aldabra as translucent with a yellowish sheen and dark 
markings. The eggs and young stages of this species are unknown. 

Measurements and proportions of representative adults are given in Table XVIII, 
and a summary for the specimens examined in Table XIX. 



Table XVIII. — Encheliophis (Jordanicus) gracilis (Bleeker). 
Measurements and Proportions of Adult Specimens. 



Collection 

Number 

Locality 



British Museum. Danish Museum. 

1861.11.28.51. . 1875. 10.5.56. . — 

Banda. . Samoa. . Ghardaqa. 



Total length . 
Length of head 

Maximum depth of head 

Maximum width of head 

Length of snout 

Horizontal diameter of eye 

Vertical diameter of eye 

Interorbital width 

Length of maxilla 
Length of pectoral fin 
Maximum depth of body 
Preanal length 



mm. 


mm. 


mm. 


. 116* 


223 


!53 


. 12-5 (11% TL) 


23 (10% TL) 


15 (10% TL) 


. 5 -5(44%HL) 


10 (43% HL) 


6 (40% HL) 


. 5 (40% HL) 


9 (39% HL) 


5 (33% HL) 


. 2. 5 (20%HL) 


4 (17% HL) 


3 (20% HL) 


. 2. 5 (20%HL) 


4 (17% HL) 


3 (20% HL) 


. 2 (16% HL) 


3-5 (15% HL) 


2-5 (17% HL) 


. 2. 5 (20%HL) 


4 (17% HL) 


3.5 (23% HL) 


• 7 (56% HL) 


10 -5 (46% HL) 


8 (53% HL) 


. 4 (32% HL) 


9 (39% HL) 


5 (33% HL) 


■ 6 (48% HL) 


11 (48% HL) 


9 (60% HL) 


. 12 (10% TL) 


21 (9% TL) 


14 (9% TL) 



* Type specimen. 
TL = total length ; HL = length of head. 



SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 301 

Table XIX. — Encheliophis (Jordanicus) gracilis (Bleeker). Summary of Principal 
Measurements and Proportions of Adults Examined. 



Variate. 

Total length 

Length of head . 

Depth of head . 

Width of head . 

Pectoral length . 

Preanal length . 



Length of head (% TL) 
Depth of head (% HL) 
Width of head (% HL) 
Pectoral length (% HL) 
Preanal length (% TL) 

N = number of specimens ; R = range of variate 
HL = length of head. 

This species occurs in the Pacific and Indian Oceans and has been recorded from 
Celebes, Indonesia, New Guinea, Cocos Islands, Pelew Islands, Fiji, Samoa, Tonga, 
Hawaii, Solomon Islands, Mozambique, Natal and Aldabra. The Pacific Ocean 
specimens have been found mainly in the starfish Culcita discoidea, while the specimen 
taken at Mozambique was in Holothuria scabra. 

Fierasfer umbratilis has long been accepted as a synonym of E. gracilis and F. 
frantii has recently also been attributed to this species (de Beaufort & Chapman, 
195 1), since there is nothing in the very careful description to enable the two to be 
separated. The species long ago recorded from Mozambique as F. punctatus must 
belong to Encheliophis for its large size shows that it was adult, yet the teeth of jaws 
and palatines were in single rows only. A fierasfer in the Danish collection, obtained 
at Ghardaqa, Red Sea, is clearly referable to F. neglectus yet does not differ sufficiently 
from the available specimens of E. gracilis to be regarded as specifically distinct. 





R. 




M. 


N. 


mm. 




mm. 


11 


116-223 




178-2 


11 


12-5-23 




18-5 


11 


5-5-*° 




7-9 


10 


5-9 




6-9 


11 


4-9 




6.5 


8 


12-21 

% 




18.3 

% 


11 


9 -6-1 1 


8 


10-4 


11 


37*5-47 


■6 


42-5 


10 


31-3-42 


9 


37'7 


11 


31-3-4° 


•0 


34*7 


8 


9-1-11 


■8 


9'8 


; M = mean value of variate ; 


TL = total length 



Encheliophis (Jordanicus) sagamianus (Tanaka), 1908 

Carapus sagamianus Tanaka, 1908, Annot. zool. jap. 7 : 40. Tanaka, 191 1, Figures and Descrip- 
tions of the Fishes of Japan : 25. Tanaka, 1927, in Figuraro de Japanaj Bestoj. Yosii, 1928. 
Annot. zool. jap. 11 : 339. 

Carapus sagamius, Franz, 1910, Abhandl. Bayer. Akad. Wiss. Suppl. 4 : 31. 

Jordanicus sagamianus, Matsubara, 1953, Jap. J. Ichthyol. 3 : 30. 

The type was found in a holothurian taken in Sagami Bay, Japan, and placed 
in the Zoological Museum of the Imperial Science College, Tokyo, no. 1751. No 
specimens have been available for study and the following description is based upon 
those published by Tanaka. 

Specific characters. Greatest recorded length 190 mm.; length of head one- 
tenth of total length ; depth of head slightly greater than width ; horizontal 
diameter of eye equal to length of snout and to interorbital width ; mouth nearly 



302 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

horizontal ; maxilla extends only to posterior edge of orbit ; vomerine teeth in 
narrow band of four rows ; pectoral fins one-third length of head ; colour in life 
unknown. 

E. sagamianus has been recorded only from Japan, at Sagami, Boshuu, Urugu 
and Misaki. It has been taken in the intestine of Holothuria monacaria and once in 
the starfish Nardoa semiregularis var. japonica. 

Smith (1955) includes this species in the synonomy of Encheliophis gracilis, but 
gives no reason why the two species should be united. 



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3 o6 SYSTEMATIC REVISION OF THE TELEOST FAMILY CARAPIDAE 

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A. 

o. 




VARIATION IN THE 
WESTERN ZOSTEROPIDAE 

(AVES) 



R. E. MOREAU 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 4 No. 7 

LONDON: 1957 



VARIATION IN THE WESTERN ZOSTEROPIDAE 

(AVES) 



BY 

R. E. MOREAU 

Edward Grey Institute, Dept. Zoological Field Studies, Oxford 



Pp. 309-433; 11 Text-figures; 14 Maps 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ZOOLOGY Vol. 4 No. 7 

LONDON: 1957 



THE BULLETIN OF THE BRITISH MUSEUM 
(NATURAL HISTORY), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical Series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

This paper is Vol. 4, No. 7 of the Zoological series. 




PRINTED BY ORDER OF THE TRUSTEES OF 
THE BRITISH MUSEUM 

Issued February 1957. Price Thirty-Five Shillings 



VARIATION IN THE WESTERN ZOSTEROPIDAE 

(AVES) 

By R. E. MOREAU 



CONTENTS 

Page 

Part i. Introduction ......... 312 

Acknowledgments . . . . . . . . • 315 

Characters of the Zosteropidae . . . . . -315 

Part 2. Colour, Pattern, Pigment and Environment . . -319 

Part 3. Dimensions and their Correlations ..... 324 

Wing-length with Altitude and Temperature . . . 328 

Tail-length with Wing-length, etc. ..... 335 

Beak-length with Wing-length, etc. ..... 339 

Dimensional Correlations and Taxonomy ..... 341 

Part 4. Continental Populations, Characters and Intergradations 342 

Introduction .......... 342 

North-east Africa ......... 343 

Remainder of Northern Tropical Africa ..... 350 

Kenya and Northern Tanganyika ...... 357 

Central Africa .......... 365 

Southern Tropical Africa ........ 368 

South Africa .......... 375 

Part 5. Insular Populations ...... 384 

Gulf of Guinea ......... 384 

Indian Ocean .......... 391 

Part 6. Synthesis .......... 404 

Summary 408 

Appendix i. Miscellaneous Notes ...... 409 

Appendix 2. Ancillary Characters . . . . . . 417 

Appendix 3. Dimensions, etc., of Continental and Insular Popula- 
tions ........... 420 

References 430 

ZOOL. 4, 7. 21 



312 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

PART 1 
INTRODUCTION 

The Zosteropidae (White-eyes) are a tropical passerine family, with a range over 
the whole of the Ethiopian Region and eastwards, through the islands of the Indian 
Ocean and India to Japan, the central Pacific and Australasia. There is however, 
a gap of over a thousand miles, presumably imposed by ecological conditions, 
between about 50 E. and 66° E., across eastern Arabia, Persia and Baluchistan. 

The family is for the most part very homogeneous, and has long been recognized 
as presenting special difficulties for the taxonomist. Stresemann (1931, 1939) has 
reviewed the populations from India eastwards, comprising about 180 forms, most 
of which are unquestionably to be placed in the genus Zoster ops. For the remaining 
Zosteropidae, those of Africa and the Indian Ocean, no conspectus is available except 
the list incidental to the Sy sterna Avium Aethiopicarum of Sclater (1930). He 
provisionally accepted 57 forms in the area and grouped them in 18 species of 
Zosterops (10 of them monotypic), 4 monotypic species of Speirops and 3 of 
Malacirops. In footnotes he mentioned 15 forms as probable synonyms ; and since 
he wrote 15 more forms have been described. More recently van Someren (1939), 
Grant and Mackworth-Praed (1943) and Mackworth-Praed and Grant (1945-46) 
have reviewed sections of the African continental birds, Bannerman (1948) has 
dealt incidentally with the West African forms, and Chapin (1954) with those found 
in the Belgian Congo. Between the various authors named there is great diversity 
of opinion about the number of forms that should be recognized and about their 
grouping into species. For example, of the n forms common to the discussions of 
both Chapin and Grant & Mackworth-Praed, Chapin admitted 4 that the other 
authors synonymized and differed from them about the specific allocation of 3 
others. Agreement on local variation (at the subspecific level) is made more difficult 
because, as I believe, changes are liable to take place in the yellows and greens of 
Zosterops skins in a short time (see examples in Note 1 of Appendix 1). It may be 
added that most of the information about the biology of Zosteropidae is merely 
casual and incidental. 

This study began as an attempt to revise the classification of the African Zostero- 
pidae but as the work progressed I became increasingly impressed with the problems 
of their variation (as distinct from nomenclature) and their correlation with environ- 
mental factors. The opportunities for investigating these correlations are 
particularly good because Africa is so mountainous and otherwise provides so great 
a variety of habitats through which the Zosterops range. 

Taxonomically, the great difficulty in this family is to determine the limits of the 
species. The Zosterops of Africa have more than once been cited as providing 
special problems when the concepts of allopatry and sympatry are applied. Lowland 
forms surround highland forms in such a way that their geographical ranges overlap 
in some cases completely, but their ecological ranges little if at all. In such cases 
the spatial barrier is, at most, of the smallest, a matter of a few hundred yards, 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 313 

and cases (detailed in the appropriate sections below) have come to light in which, 
at least locally and temporarily, none exists. How far the breeding-seasons of the 
highland and lowland forms overlap in any one locality is unfortunately not known 
for any except the special case of Nyasaland (where they do — Benson, 1953). But 
on the whole it appears that where two Zosterops occur in Africa in the same small 
area, while they may be " genetically sympatric " (Cain, 1954) only marginally, they 
cannot be regarded as " genetically allopatric " to such an extent as to exclude the 
possibility of frequent interbreeding. In such circumstances, the absence or rarity 
of hybrids, whether of the primary type (evenly intergrading) or the secondary type 
(widely and sporadically variable), can be taken to mean that the forms concerned 
are genetically so incompatible that they are best treated as belonging to different 
species. 

The subject of the present study is difficult to expound, partly because of the 
unavoidable amount of geographical and other detail, but also because of the 
difficulties inseparable from the use of trinomial nomenclature, which have been 
discussed by several recent workers (for example, Lack, 1946 ; Ellerman et al., 
1953 ; Wilson and Brown, 1953 ; Sibley, 1954). Nobody dealing with a large body 
of material from a continental area can fail to be impressed by the virtual 
impossibility of allocating every population, let alone every specimen, to a recogniz- 
able subspecies ; and if one is forced to attempt it for purposes of a check list it can 
only be at the cost of obscuring biological realities. An excellent example from the 
area under review is provided by the Zosterops of Madagascar. Originally they were 
treated as all belonging to one form ; now the palest birds and the largest birds have 
been separately named, though it has to be conceded that most of the Zosterops on 
the island are intermediate between the three subspecific types. I personally 
am deeply impressed not only with the shortcomings of the trinomial system but 
also by the harm that has been done to ornithology by the extremes to which the 
naming of local populations has been carried. Nevertheless, I regard the retention 
of some trinomials as essential, at least as a clerical convenience, especially when 
dealing with birds of a continental area. This is the spirit in which the trinomial 
is used in the present study. 

In the pages that follow I shall first discuss the probable taxonomic significance of 
colour, which has hitherto been the sole character on which the specific classification 
of the Zosteropidae of Africa and the neighbouring islands has been based, and then 
the correlations of dimensions with each other and with environmental factors. 
This will bring into prominence certain basic principles, will show the extent of 
local variation and adaptation, and will lead to the conclusion that here dimensional 
characters can be only exceptionally and to a minor extent an aid to classification. 

As regards classification at the specific level, I believe that reliance on colour 
characters has led to misleading results in the past ; and that some of the birds which 
differ conspicuously to the eye are more closely related than some of those which 
are most alike. In several islands on both sides of Africa two fully sympatric 
species of Zosteropidae occur — doubtless as a result of double invasion. By contrast, 
throughout the continent of Africa there are only two places, the upper slopes of 



314 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES 



Cameroon Mt. and the south-western Transvaal, where two different forms are fully 
sympatric. In a third area, Zululand and its neighbourhood, two forms inter- 
digitate, separated ecologically and apparently interbreeding little, while throughout 
north-eastern Africa a series of lowland forms, which I believe to form one polytypic 
species, surrounds a galaxy of isolated and well-differentiated montane populations. 
Besides the aberrant bird on the top of Cameroon Mt. which I keep in the genus 




SENECAIENSIS 



VIRENS. 



PALLIDA 



• • • MONTANE SEMEGALENSIS 

• • • POPULATIONS ISOLATED AMONC 

• • • LOWLAND ABVSSINICA. 

ZZZ ABYSSINICA. 



Map i. The ranges of Zosterops species. 



Speirops (see discussion in Part 5) I provisionally group the African Zosterops as 

shown in Map 1 : 

Z. pallida, monotypic in south-western Africa and overlapping — 
Z. virens, occupying the rest of South Africa and interdigitating with — 
Z. senegalensis , a highly polytypic species occupying most of tropical Africa and 
becoming purely montane in the northeast, where it is surrounded by — 
Z. abyssinica, a polytypic species characteristic of the dry lowlands. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 315 

These conclusions depend on a number of decisions that are by no means easy to 
take and the evidence needs to be set out in some detail (Part 4) . 

Following the description of the continental position, the insular populations on 
the opposite sides of Africa will be described. One special point of interest here is 
the evidence for double invasion, into one island after another, the evolutionary 
consequences of which can be fully appreciated only in the light of the continental 
situation. Further, the lines on which evolution has taken place in the Atlantic 
islands (Gulf of Guinea) have differed strikingly from those in the Indian Ocean 
Islands. 

ACKNOWLEDGMENTS 

This study is based primarily on the collection in the British Museum (Natural History) and was very 
greatly facilitated by the constant kindliness and co-operation of Mr. J. D. Macdonald and his staff, 
especially in handling the numerous specimens lent by other museums. 

For acceding to my requests for loans, which were often made on the most generous scale, I have to 
thank the curators of the following museums: Berlin, Brussels (Institut Royal des Sciences Naturelles), 
Bremen, Bulawayo, Chaux-de-Fonds, Chicago, Cleveland, Dakar, Durban, Edinburgh, Frankfurt a. M., 
King William's Town, Hamburg, Harvard, Leiden, Liverpool, Lourenco Marquez, Malmo, Nairobi, New 
York (American Museum of Natural History), Paris, Philadelphia. Pittsburgh, Pretoria, Stockholm, 
Stuttgart, Tervuren (Musee du Congo Beige) and Washington (Smithsonian). 

A great many individuals have helped me by replying to queries on particular points and/or by 
obtaining specimens from critical localities. It is difficult and invidious to make distinctions, but I feel 
I must in this connection give special thanks to Dr. Dean Amadon, C. W. Benson, Dr. J. P. Chapin, 
P. A. Clancey, Dr. R. M. Harwin, Dr. A. L. Rand, C. J. Skead, Dr. W. Serle, Prof. E. Stresemann, 
C. M. N. White and J. G. Williams, most of whom have also read sections of the paper in draft. Others 
to whom I am indebted in similar ways are : — 

Dr. Salim Ali, Th. Andersen, John Barlee, Dr. W. J. Beecher, Sir Charles Belcher, Prof. J. Berlioz, 
F. N. Betts, F. C. Bromley, Miss M. Courtenay-Latimer, Major R. E. Cheesman, Dr. G. Diesselhorst, Dr. 
W. N. Edwards, Dr. W. J. Eggeling, Dr. M. Eisentraut, Dr. W. A. Fairbairn, A. R. Foulkes-Roberts, Dr. H. 
Friedmann, N. R. Fuggles-Couchman, Dr. E. L. Gill, R. C. Glanville, Dr. A. J. Good, Capt. C. H. B. 
Grant, Dr. J. C. Greenway, K. M. Guichard, Mrs. B. P. Hall, Dr. F. L. Hendrickx, Dr. J. Hewitt, W. 
Hoesch, F. C. Holman, M. P. S. Irwin, Miss M. Jellicoe, Dr. G. C. A. Junge, D. W. Lamm, H. Lamprey, 
J. Last, A. Loveridge, H. von Maltzahn, Dr. A. J. Marshall, S. Marchant, Dr. G. F. Mees, Col. Ph. Milon, 

C. M. Morrison, M. E. W. North, Major W. W. A. Phillips, Dr. A. A. da Rosa Pinto, Capt. C. R. S. 
Pitman, D. C. Plowes, Dr. G. Popov, Dr. H. Poisson, Dr. A. Prigogine, J. F. Pringle, Mrs. M. K. Rowan, 
E. G. Rowe, Dr. G. Rudebeck, Dr. E. Schelpe, Dr. H. Schouteden, Dr. H. Scott, K. D. Smith, R. H. 
Smithers, R. Stowell, Dr. C. Vaurie, Dr. R. Verheyen, L. D. E. F. Vesey-Fitzgerald, J. Vincent, Dr. J. 
Vinson, R. Wagstaffe, A. S. Watt, C. Whybrow, Dr. J. M. Winterbottom, C. G. Young, Dr. G. Zink. 

Treatment of certain specialist points would have been impossible without expert assistance, from Dr. 
L. Auber, of the Wool Industries Research Station, Leeds, in connection with pigmentation, from Mr. 
N. J. T. Bailey, Lecturer on the Design and Analysis of Scientific Experiment, Oxford, who arranged for the 
statistical analysis, and from the Library of the Meteorological Office, Air Ministry, which recommended 
and lent a mass of climatological records. These were kindly supplemented by information given by 
various authorities in Africa, especially those of Angola and the Union of South Africa. For work on 
the maps and figures I have to thank Mrs. R. Heelas, Miss P. M. Moreau and Miss M. Potter. 

Finally, I have benefited greatly from discussions with Dr. A. J. Cain, Dr. D. Lack and Dr. H. W. 
Parker, who have read the whole manuscript in draft, and with Dr. E. Mayr, Dr. C. G. Sibley and Dr. 

D. W. Snow. 

SOME CHARACTERS OF THE ZOSTEROPIDAE 
AND OF THE AFRICAN MEMBERS 

In appearance the Zosteropidae are rather " ordinary " passerines nearly all less 
than 14 cm. long. The size range is not great ; omitting two aberrant insular forms 
usually kept in the genus Speirops, within the present area the following are the 
extreme measurements : wings 49-69 mm., tails 30-53, beaks 11-17. Morpho- 
logical characteristics of the family are absence or extreme reduction of the outermost 



3 i6 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

primary, a fimbriated bifid tongue and a ring of silky-white feathers round the eye. 
The Zosteropidae are all short-winged birds, of direct flight, and thoroughly arboreal 
habits. The beak is rather sharp and slightly decurved. 

Certain behaviour-characters of the family are significant in any consideration of 
their evolution : 

(i) The birds have a wide range of food, taking fruit, insects and nectar, 
and in this respect there seems to be no specialization, local or other. Their 
feeding habits often bring them into association with sunbirds, alongside which 
they are always placed in taxonomic order. 

(2) Individuals of most forms have a strong tendency to flock. Consequently 
birds journeying outside their normal area are likely to do so in groups and 
successful colonization is thus facilitated. 

(3) White-eyes are liable to erupt and cross several hundred miles of sea. 
They have colonized more oceanic islands than any other passerine genus. 
That the Australian Z. lateralis invaded New Zealand across more than 1,000 
miles of sea is a matter of history ; and that Norfolk Island, some 500 miles 
from any other Zoster ops station, has been successfully colonized three times is 
certain from its three different forms of Zoster ops. 

(4) In conflict with this, the presence of highly differentiated populations 
on both marine islands and continental mountains within sight of each other 
seems evidence that the birds of some populations must be profoundly sedentary. 
An extreme instance is that of the Z. rendovae group (cf. Mayr 1947), where 
water-gaps of less than two miles separate subspecies. 

Especially in a continental area, such as Africa, the birds would be expected to be 
most sedentary in the most equable environment (evergreen conditions) and least 
sedentary where the seasonal changes are greatest. There is good indirect evidence 
of this : 

(a) A series of populations (52-54 in Appendix 3) having identical plumage 
and living in an evergreen-forest climate on the eastern rim of the Congo basin 
has average wing-length 61-3 mm. at 7,000 ft., 55-9 mm. forty miles to the west, 
at 4,300 ft., and 53-1 mm. twenty miles west again, at 3,000 ft. 

(b) In East Africa very distinct populations live on forested mountains, 
separated from each other by as little as twenty miles. 

(c) By contrast, the Zoster ops ranging through the belt of deciduous thorn- 
bush south of the Sahara has not differentiated to the subspecific level in the 
three thousand miles from Senegal to the eastern Sudan. 

There is little direct evidence of movement : the most definite is that the 
Zosterops characteristic of the very dry country of southern South West Africa 
sometimes appear for a few days in Windhoek (W. Hoesch in litt.), some 150 miles 
from where birds of this kind are known to be resident. 

Birds usually assigned to the genus Zosterops comprise more than four-fifths of 
the Zosteropidae. Nearly all of these have the upper parts more or less green — 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 317 

with a range of colour from dull pale grey-green to bright yellow-green and rich 
olive-green ; their throats and vents are more or less yellow. They have no distinc- 
tive patterns on wings or tails and no spots or speckles anywhere. There is no 
sexual dimorphism except that females tend to be a little smaller and, like juveniles, 
a little duller in plumage (having less carotenoid). Individual variation is, however, 
high — so much so that I should expect the results obtained by Marples (1945) to 
find a parallel in African populations if comparable series were examined. He was 
able to classify the belly-colour of Z. lateralis, trapped at Dunedin, New Zealand, in 
winter into nine different categories. 

Apart from differences in shade and in dimensions, the main variations between 
populations are in three features : 

(1) The width of the ring of white feathers round the eye (a feature that 
unfortunately does not lend itself to accurate measurement). Since an eye- 
ring exists in the vast majority of the Zosteropidae, and often is retained in 
aberrant forms whose other characters are much modified, it might be supposed 
to be of biological importance. But there is no evidence of its function, whether 
in display or in any other way, and within Africa it varies in extent from a 
conspicuous white patch, covering half the side of the head, to an almost 
imperceptible rim round the eye. Moreover it is sometimes altogether lost 
in otherwise " normal " Zosterops. On the whole it cannot be regarded as 
diagnostically useful except at the subspecific level. 

(2) Markings on the forepart of the head. These are usually limited to lines 
above the lores and/or areas on the forehead where the melanin is reduced, so 
that they appear more yellow than the rest of the upper parts. If these brighter 
areas were sharply demarcated, they would be more comparable for taxonomic 
purposes. 

(3) The underparts between the upper breast and the under tail-coverts. 
These are nearly always either yellow, with some green wash, especially on the 
flanks, or whitish washed with greyish or brownish. This feature is discussed 
specially in Part 2. 

Particularly within the Zosteropidae discussed in this paper, there are extremely 
few clear-cut differences in pattern. Nearly all those perceptible between popula- 
tions are merely a matter of degree — nuances of intensity of melanin or carotenoid ; 
while the one striking difference in pattern in the continental African birds, yellow 
or no yellow on the belly, which has been accepted as a specific difference, cannot be 
regarded as a reliable guide to relationships, as shown in Part 2. 

Since the genus Zosterops includes over 200 recognized forms and for the most part 
varies only within the narrow limits indicated above, it is easy for widely separated 
populations to show resemblances that must be the result of convergent evolution 
rather than of close affinity ; and the systematise s task is correspondingly more 
difficult. For example Madagascar Zosterops are very like those of New Guinea 
(among others), those of the Australian mangroves (lutea) like those of the East 
African lowlands {flavilateralis) , and those of Annobon Island in the Gulf of Guinea 



3 i8 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

(griseovirescens) like those of Christmas Island in the Indian Ocean (natalis). Again, 
within our area, Madagascar Zosterops differ from those of south-western Abyssinia 
only in lacking golden foreheads and in having a little more melanin generally 
(characters that are not independent). Moreover, this difference is comparable to 
that (also confined to colour) between Zosterops from Madagascar and from India ; 
for, while most of the latter are much yellower than the former, Nilgiri birds are 
intermediate. Again, within Africa itself, some Zosterops, separated by hundreds of 
miles occupied by other forms, are indistinguishable, such as those of the mountains 
on the Sudan-Uganda border and some of those in Tanganyika Territory. 

In such circumstances, where the appearance of the birds can be so unsafe a guide 
to their affinities, as many ancillary characters as possible must be considered. 
Among the most important of these is habitat preference. In his study of the 
eastern Zosteropidae Stresemann (1931) regarded their ecology (particularly their 
preference for lowland and highland habitats respectively) as an important guide 
to their relationships at the specific level. (Yet he was forced to postulate consider- 
able change in ecological preferences in arriving at his grouping of forms into the 
polytypic Z. citrinella and still more in his enlarged concept of Z. palpebrosa 
(Stresemann, 1939)). There is no doubt that for African birds in general habitat 
preferences are highly specific, and in some cases generic, especially as between 
forest and non-forest and between highland and lowland habitats (cf . the analysis in 
Moreau, 1954). Unfortunately in the African Zosterops these distinctions and 
preferences are less clear-cut. 

This is presumably due in part to their way of life : they are not ground-feeding 
birds, which in evergreen forests are immersed in a specialized eco-climate, nor 
forest-canopy birds, which may be limited by the distribution of certain species of 
fruiting trees or of tree-holes. Instead, the Zosterops have a wide range of diet and 
no specialized nesting-site ; moreover those forms which are attracted to evergreen 
forest appear to belong to the edges rather than the depths. (There is no Zosterops 
at all in the main Congo forest area, even in the clearings.) Also, most forms of 
Zosterops, including those of evergreen forest, seem to adapt themselves readily 
to whatever man-made conditions provide trees and associated food. Even those 
Zosterops which are always cited as forest birds, such as those in the southern 
Cameroons and on the East African mountains, may perhaps be looked upon as 
birds that use the arboreal associations dominating the locality rather than birds 
ineluctably dependent on forest, as most members of the evergreen-forest 
communities seem to be. If this view is correct, then different populations of the 
same species of Zosterops may be found occupying in one part of Africa savanna trees 
or even thorn-bush, and in others evergreen forest, with more or less ecotypical 
modification of characters. As will be pointed out, this is what happens in 
Madagascar, where Zosterops unquestionably of the same species occupy all types 
of vegetation and climate from the " subdesert " of the south-west to the east 
coast with over 100 inches of rain. Again, in South Africa birds that cannot be 
separated taxonomically are found breeding in a range of climate and natural 
vegetation from the coast of the Indian Ocean and associated evergreen forests, to 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 319 

the " high- veldt " of the Transvaal with its far greater daily and annual range of 
temperature and far drier and more deciduous " bush ". This is not to deny that 
such birds as the montane kikuyuensis of the eastern Kenya highlands and the low- 
land flavilateralis are usually separated ecologically — even though in a marginal 
locality, such as near Nairobi, they may frequent the same garden. But it is at 
least clear that, in the key problem of which forms of the African Zoster ops should 
be allocated to which species, in cases of difficulty the habitat preferences are not 
sure guides. 

Other ancillary characters which have been considered are (1) colour of beak, iris 
and legs, (2) form of beak and tongue, (3) wing-formula, (4) voice, (5) feathering 
sequence of nestling. Certain of these, especially (2) are probably particularly 
adaptive and labile, but in any event, on examination, none of them seems to be 
capable of giving important clues in difficult cases. Details are therefore relegated 
to Appendix 2. Only two points in connection with them need be noted here : 

(a) Such differences in wing-formula as exist seem to have neither taxonomic 
nor consistent ecological relationships. Two of the forms with the bluntest 
wings belong to dry country, but in two others of equally dry country (with open 
thorny trees) the character is not so marked ; and on other grounds there is 
little doubt that the four forms belong to at least three different species. 

(b) Beaks and tongues are much alike in all the continental African and nearly 
all the insular birds under consideration, irrespective of habitat. 

(c) The songs of some of the most different-looking African Zoster ops, which 
have usually been allocated to different species, are apparently alike. 

In attempting to discuss relationships and evolution one further difficulty is 
encountered, namely, apart from certain details of the plumage (see Part 2), there is 
no sure indication in the Zosteropidae of which features are the most readily modified. 
For example, albogularis of Norfolk Island has attained giant stature while retaining 
the presumed ancestral colouring, including full carotenoid and white eye-ring. Yet 
in a few forms the eye-ring seems to have been one of the first characters to be 
reduced or lost (at the subspecific level in uropygialis of one of the Kei Islands) . On 
the other hand, one of the Seychelles birds, modesta, has lost all its yellow pigment, 
and become uniform grey, without becoming abnormal in size, proportions, beak or 
eye-ring. In the insular Zosteropidae, in fact, it seems that a rather limited variety 
of evolutionary changes can take place, with one exception, in almost any order. 

part 2 

COLOUR, PATTERN, PIGMENT AND ENVIRONMENT 

As indicated in Part 1, the most prominent colour differences between forms of 
Zosteropidae is that some have yellow pigment (more or less washed with green) on 
the belly and some do not. Stresemann (1931) noted in discussing the Indo-Pacific 
Zosteropidae that absence of yellow from the belly is much commoner than its 
absence from either the throat or the under tail-coverts. In fact, this last occurs 



320 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 



only if yellow pigment is lacking from the whole of the rest of the underparts — 
which usually means from the upper parts as well. These generalizations apply 
equally to the African and insular Zosteropidae dealt with in the present study. It 
may be noted also that absence of yellow from the belly is unknown as an individual 
abnormality in a yellow-bellied population, though absence or reduction of carotenoid 




Map 2. The geographical distribution of Zosterops belly-colour. 



from the whole plumage occurs, as well as loss from defined asymmetrical patches 
(see Note 2, Appendix i). 

All who have dealt with the taxonomy of the African Zosterops have regarded the 
populations that have yellow bellies as specifically distinct from those which have not, 
though there has been no agreement about how many species of either type should 
be recognized. By contrast, however, in a number of polytypic Indo-Pacific 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 321 

Zosterops species Stresemann (1931) and Mayr (1944) include both populations with 
yellow on the belly and populations without. Stresemann concluded that the 
difference probably depends on a single genetical factor. This must be true in any 
case where the two types co-exist and are proved to interbreed without producing 
any birds of intermediate plumage, but this condition does not seem to have been 
completely satisfied anywhere and at least in Java the evidence is against a single- 
factor explanation (Mees, 1951 ; Note 3, Appendix 1). There is in the island a 
complete range of variation between the yellow-bellied gallio and the grey-bellied 
buxtoni ; and it is instructive to note that many of the specimens are so like one of 
the parents that they would have escaped detection as hybrids if they had been seen 
in isolation. 

In Africa what has been discovered in recent years about the relationship between 
the two types of Zosterops makes it virtually certain that here also belly-colour is 
not a specific character. For one thing, the grey-bellies occur only in widely 
separated areas (Map 2) , isolated from each other by yellow-bellied populations that 
occupy equivalent niches in the intervening country (for details of distribution see 
Part 4). 

The grey-belly locations fall into three groups : 

North-east Africa. 

In the Abyssinian highlands grey-bellied poliogastra and yellow-bellied 
kaffensis are almost allopatric and no intermediates are known. 

The lowlands are occupied by grey-bellied birds (abyssinica and omoensis) 
except in the west-centre and the south-east. They are allopatric to yellow- 
bellied birds except near Lake Tana. 

Kenya and Tanganyika Territory. 

Here the numerous isolated mountains form an ecological archipelago. 
Grey-bellied Zosterops occupy exclusively (1) Kulal Mt. (kulalensis) , (2) the 
Teita group (silvana), (3) the South Pare range (winifredae) . The intervening 
and other mountains are occupied exclusively by various forms of yellow- 
bellied Zosterops, and each of the three grey-bellies is more like its nearest 
montane yellow-belly neighbour than any other. 

South Africa. 

In the Cape Province grey-bellies predominate in the west, yellow-bellies 
(otherwise identical) in the east. They overlap considerably and in the middle 
they are proved to interbreed commonly. Unfortunately the data do not 
suffice to settle the genetics. 

Another form lacking yellow on the belly occupies South West Africa and 
overlaps the yellow-belly marginally, but in this case there is no evidence of 
interbreeding. 

The foregoing data, taken as a whole, certainly suggest that in Africa colour of 
belly is a character with a simple genetical basis and that grey-bellied populations have 
developed repeatedly. In East Africa the montane grey-bellied forms have every 



322 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

appearance of having arisen as so many different mutants from the neighbouring 
yellow-bellies. It is concluded that in the Zosterops of Africa, as in those of the East, 
belly-colour is not necessarily a specific character. There is nothing to indicate how 
the present geographical pattern arose. It is difficult to believe that the character 
" grey-belly " is directly adaptive and, even with personal knowledge of East 
African mountains, I cannot suggest what factors can have operated particularly on 
Kulal, Teita and South Pare to select any characters with which " grey-belly " 
might be linked. It may be added that, though probably the Kulal and Teita birds 
are hardly to be numbered in thousands, the populations are far too big for the 
Sewall- Wright effect to be accepted as bringing about the present situation. 

Much of the taxonomic discussion about the African continental Zosteropidae has 
turned upon the exact shade of yellowish green or greenish yellow of the upper parts, 
and the amount of green on the sides of yellow underparts. The colour intensity 
of the blackish lores and of the remiges and rectrices has also been adduced as a 
character. These are, however, not independent features because they are all 
correlated with the nature of the melanin present in the plumage as a whole and with 
the degree of its concentration. If much melanin is present in the upper parts, the 
yellow on the front of the head is, as a rule, less than in closely similar populations 
that have the upper parts as a whole not so dark. Conversely, the amount of greenish 
wash on the flanks generally varies with the darkness of the upper parts. 

The physical basis of the coloration of the Zosteropidae does not seem to have been 
described anywhere and I am greatly indebted to Dr. L. Auber of the Wool Industries 
Research Station at Leeds for making microscopical investigations for me. It is 
hoped to extend and publish the results separately. Dr. Auber rinds that in 
Zosterops pigments of only two groups are present, carotenoid (yellow) and melanin 
(three types). In the terminal (exposed) part of each contour feather the barb 
and /or the individual barbules have more or less of their length yellow and the rest 
melanic. Superimposed on each other in the plumage, they give, by a " lattice " 
effect, green. The exact shade depends on (a) the relative extension of the yellow and 
the melanized parts of each barbule, (b) the shade of each of the pigments, (c) their 
concentration and distribution within the tissue. 1 This last is especially important 
with the carotenoid, for the intensity of the yellow colour depends mainly on the 
thickness of the layer containing the pigment. This is especially important in the 
barbs, where the thicker the cortex (the layer holding the carotenoid) the more 
intense the yellow, towards golden or even orange. This effect appears to be mainly 
responsible for the striking difference between the pale yellow on the forehead of 
abyssinica and the strong reddish-golden of the Kenya kikuynensis. In the former 
the unpigmented central core (medulla) of the barbs is wide, while in the latter it is 
so reduced that the barb forms practically a column saturated with carotenoid. 
Here, then, is a case in which the colour depends on the anatomy of the feather 
as well as on the pigment. 

1 The conventional phrase " grey wash " or " green wash " is misleading in so far as it implies a 
pigment applied externally. In the Zosterops an impression of " grey wash " is given in an area of 
plumage that contains some melanin but little or no yellow pigment, and of " green wash " by an increase 
in melanization in the presence of yellow pigment. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 323 

With the melanins the optical effect depends largely, but by no means wholly, on 
the type of melanization. In approximate agreement with the observations of 
Gornitz (1923) and Frank (1939) on other groups of birds, Dr. Auber distinguishes 
three types of melanization in the African Zoster ops, namely : 

(A) Rodlets more than ifi long, coated with blackish pigment and arranged 
with their axes parallel to the main axis of the barbule. 

(B) Granules shorter and more irregular than (A) and which under the 
microscope look more brownish. 

(C) Very small irregular granules, bright reddish brown. This type is rarer 
than the others in the African Zosterops ; it predominates (and causes a reddish 
colouring) only in the flank-feathers of the dry-country birds of south-western 
Africa (pallida) and in two insular birds (semiflava and mayottensis) living in a 
much damper oceanic climate. 

Throughout Africa the melanin in the plumage of each form of Zosterops is pre- 
dominantly or (usually) exclusively, of either (A) type or (B) type. Since the 
second pigment is much browner than the first it might be supposed that the type 
of melanization would proclaim itself to the eye but, for reasons not altogether clear, 
this is not always so. The outstanding example of this is afforded by the abnormal 
birds of the Gulf of Guinea islands, which are browner than any other Zosteropidae, 
yet contain much (A) type melanin. Again, the dry-country birds of rather dingy 
appearance in both north-east and south-west Africa, abyssinica and pallida, are 
dominated by (B) type, while the Zosterops of the humid Sao Tome and Principe 
Islands, the upper parts of which look very similar, are filled with (A) type. Also, 
while in all the richer green Zosterops of Africa (A) -type melanization predominates, 
the Cape bird, which looks similar, is mainly (B) ; and such melanin as exists in the 
brightest yellow Zosterops in Africa, senegalensis of the dry belt south of the Sahara 
and anderssoni of the Rhodesias, is entirely (A) . Incidentally, this explains the full 
blackness of the loral spot in these latter birds, a feature at first sight surprising in 
a Zosterops whose general plumage gives so little sign of black pigment. 

In certain South African forms the proportion of (A) and (B) melanization in 
the individual varies with the humidity of the environment. Dr. Auber has found 
that in the birds of the western Cape Province (capensis) the admixture of (A) in 
the predominant (B) is greater in the humid Knysna area than in the arid Kamies- 
burg. Also, in virens of eastern South Africa the melanization is chiefly (A) in the 
humid Natal highlands, (B) in a dry-veldt habitat in the Transvaal. By contrast, 
senegalensis has all its melanin of (A) type, even in the driest environment (cf. a 
specimen from Maiduguri, near Lake Chad). This presumably means either that 
there is a different (undetected) environmental factor operating or, more probably, 
that senegalensis is physiologically (genetically) different from these South African 
birds and also from the other dry-country Zosterops (of north-eastern Africa), 
which are characterized by (B) melanin. 

On the whole, as will be seen from the descriptions in Part 4, the more richly 
green birds throughout Africa occur in the more humid areas and this is everywhere 



324 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

linked — by whatever mechanism — with a greater intensity of (A) melanization in 
the plumage. There are very strong indications that the change takes place clinally 
in step with change in vegetation-type. The change is less closely correlated with 
total annual rainfall because other climatic factors, such as seasonal cloud-cover, 
extent of occult precipitation, and above all distribution of rainfall through the year, 
all determine the biological effectiveness of the rain. Thus, less than 60 inches of 
rain can suffice for evergreen forest if there is no long dry season, but not otherwise. 

From all of the foregoing it will be evident that on the whole the African Zosterops 
conform to Gloger's rule, under which, as summarized by Mayr (1947), melanins 
increase in the warmer and more humid climates and in arid climates reddish- or 
yellowish-brown melanins increase at the expense of black. But in the African 
Zosterops it appears that melanins increase in the warmer climates only where the 
humidity is not reduced ; and, so far as the type of melanin is concerned, the 
exclusive presence of black in the dry-country senegalensis is an exception to Gloger's 
rule. 

To summarize this section, it appears that belly-colour cannot in these Zosterops 
be taken as a guide to taxonomic affinities, nor can general plumage-colour nor, as 
a rule, the type of melanization present — which often does not proclaim itself to the 
eye. 

PART 3 

DIMENSIONS AND THEIR CORRELATIONS 
IN CONTINENTAL ZOSTEROPS 

The inquiry into these correlations was started in the hope of finding peculiarities 
and discontinuities that might help to elucidate the taxonomy of the birds. This 
section has taken its present form thanks to the statistical analysis arranged for 
by Mr. N. J. T. Bailey in his department of Design and Analysis of Scientific Experi- 
ment, at Oxford ; and what follows here owes a great deal to discussions with him, 
with Mr. John Barlee and with Dr. H. W. Parker, who has devoted much time to the 
problems here raised. 

The variations investigated are those of wing-length, tail-length and beak-length 
in relation to temperature and altitude, and in relation to each other. Investigation 
of these environmental effects is complicated by the fact that temperatures vary 
with altitude. Diverse as the temperatures are in different parts of Africa at the 
same altitude, in any given locality the temperature falls by about 3-5° F. for every 
1,000-foot rise in altitude. The other factor that changes consistently with altitude, 
air-pressure, falls off by about 2*5% 1 of sea-level pressure for every 1,000 ft. of 
altitude and is not subject to important local variations. 

By a suitable statistical technique the relations of dimensions to environmental 
temperature can be separated from altitude effects and considered independently, 
with special references to Bergmann's rule. As summarized by Mayr (1947), this 

1 This figure is net, taking account of the elfect that the reduction in temperature has on the air- 
pressure. I am indebted to Dr. Parker for pointing out also that air-density is affected by humidity, 
being reduced as humidity increases ; but the effect is small enough to be ignored for the present purpose. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 325 

is to the effect that " smaller geographic races of a species are found in the warmer 
parts of the range, the larger-sized races in the cooler ". Under " Rules applying 
to birds only " it is also stated that " the wings of races that live in a cold climate 
or in the high mountains are relatively longer than those of the races which live in 
the lowlands or in a warm climate ". It is not clear whether the word " relatively " 
means " relative to body-size "; if it does, then it must be noted that none of the 
references quoted by Mayr in support of this statement provide conclusive evidence. 

Comparisons for this purpose have usually been made in the broadest terms, 
using latitude as an indicator of temperature, rather than actual meteorological 
data, and without discussing which elements of the environmental temperature 
are the most significant. It might be expected, a priori, that selection would be 
exercised by the more extreme conditions encountered ; Huxley (1942) in fact 
suggested that in the temperate zones these would be primarily the winter minima 
and in the sub-tropics (presumably even more in the tropics) the summer maxima. 

The term used in the formulation of Bergmann's rule is " size ", which presumably 
means the size of the animal as a whole ; and consideration of the relations between 
size and temperature turns ultimately on the fact that, other things being equal, 
the heat exchange between a body and its environment depends on the ratio between 
its volume and its surface. The bigger and /or the more compact the body, the 
less rapid (per unit of mass) its heat-exchange. The bigger the body, the better it is 
fitted to withstand cold ; and the smaller the body the more easily it can, through 
radiation and evaporation, lose heat. 

For vertebrates, weight is a useful indicator of size, but for most birds so little is 
known about the weight that in ornithology wing-length is almost always used as an 
index of body-size, with the tacit assumption that the relationship is linear. 
(Incidentally, the standard measurement of wing-length is not skeletal, but effectively 
that of the longest feather.) Now, if the shape of the wing and all the other bodily 
proportions remain constant, the " lift " provided by the wing varies as the square 
of the wing-length, while the mass of the whole bird varies as the cube. In these 
circumstances, if efficacy of flight is to be maintained it can be by an improvement 
in flying technique or by the length of the wing increasing faster than the linear 
dimensions of other parts of the body. In this case wing-length, so far from bearing 
a linear relation to the mass of a bird, would bear a complex and also continually 
changing relation to it, and hence to the bird's potentiality for heat-exchange. 

Since no weights are recorded for African Zoster ops, no data exist for testing the 
ratio of wing-length to mass in any of the populations 1 , so that a link essential for 
any comprehensive discussion of size /temperature relationships is missing. Two 
further caveats are needed. The first is that the African Zoster ops cannot all be 
claimed to belong to one species, which is a condition of Bergmann's rule ; but, as 
will be concluded, most of them appear to be conspecific. The other caveat is that 

1 An investigation of the mean weight of bird populations for statistical purposes such as the present 
would need especially long and critically collected series because the weight of the individual fluctuates 
so much with the time of day and season of the year. In Zoster ops at Dunedin, N.Z., Marples (1945) 
found that mean weights of samples trapped on different days in the course of the year varied between 
12 and 15 gm. 

ZOOL. 4, 7. 22 



326 VARIATION IN THE WESTERN ZOSTEROPIDAE (AYES) 

heat-exchange depends on insulation as well as on mass and configuration ; we have 
no precise information on this point. It can only be said that the Zoster ops of the 
higher, cooler, environments appear to have thicker plumage than the others. 

The dimensional data 

For the present study about 2,500 specimens have been measured, a large propor- 
tion of all those housed in the museums of the world. As will be seen from Appendix 
3, there are eighty continental populations of which enough specimens have been 
examined to give results useful for statistical treatment. Some of these " popula- 
tions " coincide with named forms, but many of them do not. The latter are of 
three types : (a) geographical sections of forms with very extensive ranges, for 
example the respective Northern Rhodesian, Southern Rhodesian, Nyasaland and 
S.W. Belgian Congo populations of anderssoni ; (b) altitudinal divisions, as between 
the Zosterops of the eastern Kenya highlands above and below 9,500 ft ; (c) popula- 
tions that are transitional and /or were not known to earlier workers (so that no 
subspecific name has been applied to them in the past). Other specimens and their 
dimensions are mentioned in the text as necessary. 

All the measurements given in Appendix 3 were made by me personally, so that 
any subjective error should be fairly constant. Wings have been flattened and 
straightened, so that the measurement recorded is the greatest possible. Tails 
have been measured with one point of the dividers pressed down between the two 
middle feathers. Beak measurements are particularly difficult in Zosterops because 
of the lack of abruptness in transition from culmen to skull. At first, indeed, I 
felt that this measurement could not usefully be made, but eventually I found that 
fairly satisfactory results were possible — cf. the length-ranges in Appendix 3. In 
each case the point of the dividers was slid with gentle pressure up beyond the base 
of the culmen until it was decisively checked. 

In the entire series of continental birds examined wing-length varies from 49 to 
69 mm. in individuals and from 51-3 to 64 mm. in means. Extreme variation 
within populations is nearly always 4-7 mm., i.e. up to about 12% of the mean. 
The few populations in which it is much greater are either represented by 
exceptionally long series or could possibly be sub-divided if more specimens, suitably 
distributed geographically or altitudinally, were available. An example is provided 
in the highlands west of Lake Edward (populations 48 and 49 in Appendix 3) . The 
specimens at first examined showed within this small area wing-lengths with the 
exceptional range of 54-65 mm. Later it was found that 31 birds from an average 
altitude of 4,700 ft. (the highest 5,000 ft.) measured 54-60 mm. and 10 from an average 
altitude of 6,800 ft. 58-65 mm. In general, individual variation in tail-lengths and 
beak-lengths is proportionately greater than in wing-lengths, perhaps partly because 
of the greater difficulty in making the measurements. 

A small source of bias in the figures for the means comes from the fact that males, 
females and unsexed birds have all been used, provided that they showed no 
immaturity and no moult in the feathers to be measured. It would have been more 
satisfactory to use birds of only one sex, but this was rejected for two reasons. Firstly, 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 327 

the sexing of collected specimens is not always reliable and in some populations 
specimens that are not sexed form a large part of the material available. Secondly, 
although females are on the average smaller than males in most populations, the 
difference does not exceed 2 mm. in even the biggest Zoster ops. Moreover, in nearly 
all the populations the proportion of sexed female specimens varies only between 
one quarter and one half of the total, and within these limits the most the mean 
could be biassed by this variable is about 0-5 mm. This is not enough to invalidate 
the general conclusions to be drawn later. 

The meteorological data 

For each population in Appendix 3 a mean altitude has been calculated for the 
specimens actually measured. (This mean does not necessarily agree with the 
mean altitude of the habitat of the Zosierops population concerned over its whole 
range.) Where no altitude is specified on the label of a specimen it has been estimated 
from its locality and such knowledge of the topography as could be obtained. It 
is thought that most of the mean altitudes given in Appendix 3 are accurate to 
within 500 ft. 

These mean altitudes have been used as the datum-lines for calculating the 
temperatures that are given for each population in Appendix 3, namely, the mean 
minimum of the three coldest months and the mean maximum of the three hottest. 
(From these figures approximate annual means and annual ranges can be calculated 
as needed.) Thanks to the assistance of the Meteorological Office, Air Ministry, 
and supplementary information from African meteorological services, temperature 
data have been assembled for about 1,000 stations south of the Sahara. Within 
the geographical range of most of the populations in Appendix 3 a number of temper- 
ature records are available, some of the stations being as a rule higher and some lower 
than the mean altitude of the specimens concerned. From these records mean 
temperatures have been calculated and also the mean altitude of the stations. 
Where this differs from the mean altitude of the specimens available an adjustment 
has been made at the rate of 3-5° F. per 1,000 ft. Extrapolation has been used in 
two types of case : 

(a) Where, as in birds from high altitudes in eastern Kenya and Mt. Cameroon, 
mean altitude of the specimens is thousands of feet above that of the local 
meteorological stations, the same factor of 3-5° F. per 1,000 ft. has been used. 

(b) Where, as in the case of populations very narrowly localized, e.g. on single 
isolated peaks, meteorological stations are available only elsewhere in the same 
general area, a mean has been calculated from the data of the nearest ecologically 
comparable stations (and then the altitude correction applied if necessary) . 

Especially in the latter type of case, unknown local meteorological factors may 
invalidate to some extent such extrapolation, as personal experience of East Africa 
suggests, and for two of the populations in Appendix 3 I have thought it undesirable 
to attempt extrapolation. I am aware that some of the temperature means arrived 
at are open to criticism ; but all have been calculated as objectively as possible and 



328 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

the correlations that are about to be described show that in general the methods 
employed are valid. 

WING-LENGTH IN RELATION TO ALTITUDE 
(AIR-PRESSURE) AND TEMPERATURE 

In Text-figs. 1-3 the mean wing-lengths detailed in Appendix 3 are plotted respec- 
tively against altitude, minimum temperature and maximum temperature (as defined 
previously). In some respects these diagrams are illuminating, but they can, as 
will be shown, also be thoroughly misleading, for the reason that the three environ- 
mental factors are not independent. 

In order to separate the effects of altitude and temperature, partial regression 
coefficients have been calculated, which give a measure of the influence that each 



65 r 



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ALTITUDE (FEET) 

Fig. 1. Wing-length /altitude correlations. (Black circles, South African populations ; dotted 

circles southern tropical.) 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 329 

factor would have if it varied while the others remained constant. If w = wing- 
length (in mm.), a = altitude (in thousands of feet), n = minimum temperature 
(°F.) and x = maximum (°F.) the partial regression equation is : 

w = 59-8 -j- 0-690 — o-28n -f 0-12%. 

Each of these regression coefficients is highly significant statistically, since their 
standard errors are only 0-13, 0-04 and 0-05 respectively. It is to be noted, however, 
that each cofficient represents an average, and the data do not suffice for an 
investigation of whether the relationship in each case is rectilinear or not. From 
the purely statistical point of view the equation fits the observed data so closely 
as to warrant the conclusion that no environmental factor having a comparable 



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70 « 60 55 SO 45 40 35 

MINIMUM TEMPERATURE («F) 

Fig. 2, Wing-length /minima correlations. 



33© VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

effect on the wing-length remains unconsidered. From the biological point of view 
this is in at least one respect questionable, as will appear. 

Meanwhile, subject to these comments, it is concluded from the equation that on 
the average, over Africa as a whole : 

(i) Zosterofs wings tend to increase 0-69 mm. for every 1,000 ft. of altitude. 
Since the altitude range of the combined Zoster ops habitats is 10,000 ft., the 
total potential effect of this factor is about 7 mm. (12% of the overall mean 
wing-length) . 

(2) Zosterops wings tend to decrease 2-8 mm. for each rise of io° F. in the 
minimum. With extremes of mean minima 35 and 72 F. (Appendix 3), the 
total potential effect is about 10 mm. (17% of the overall mean wing-length). 

(3) Zosterofs wings tend to increase in length by 1-2 mm. for each io° F. 



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ICO 95 90 65 BO 75 70 6S 

MAXIMUM TEMPERATURE («0 

Fig. 3. Wing-length /maxima correlations. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 331 

rise in maximum temperature of the environment. Since the lowest maximum 
in Appendix 3 is 65 F. and the highest ioo°, the total potential difference in 
wing-length attributable to this cause in Africa is only about 4 mm. (7% of 
the overall mean wing-length) . 

Wing-length and altitude 

The equation shows that altitude has an important effect on wing-length that has 
nothing to do with temperature and hence presumably operates through the reduced 
air-pressure No conclusive evidence of such an effect appears to have been 
presented hitherto for any type of bird. Long critical series of weights and wing- 
areas would be required for a closer examination of this phenomenon. 

Since changes in air-pressure purport to be a direct cause of the changes in wing- 
length, then, when changes of pressure with altitude are plotted along with the 
changes in the " lift " provided by the wing, i.e w 2 , the two " curves " might be 
expected to be of similar form. In fact they are not, because the pressure /altitude 
curve is very close to a straight line, while that of altitude \w 2 is quite different. 
The explanation may be complex. There is the purely statistical point that the 
addition of 0-69 mm. to the wing-length per 1,000 feet, is an average one ; as 
already noted, this may conceal a non-rectilinear relationship that the data are not 
" good " enough to disclose. Another, and strong, possibility is that the relation- 
ship between the wing-length and the mass of the bird is disturbed by various 
anatomical adjustments necessitated by changes in air-pressure. For example, in 
the species investigated by Norris and Williamson (1955) the ratio of heart-weight 
to total weight increased with the altitude at which the birds lived ; and similar 
changes in lung-capacity may be expected. 

Text-fig. 1 shows that no population of small Zosterops, with wing averaging 
less than 58 mm., occurs above 5,000 ft. Another prominent feature of Fig. 1 is 
the grouping of the South African populations (black circles). Their appearance 
of being long-winged for their altitudes is undoubtedly due to the fact that South 
African temperatures are lower than those of tropical Africa at the same altitudes 
(cf . Appendix 3) ; it will be seen that this segregation of the South African points 
is not repeated when the wing-lengths are plotted against the minimum temperatures 
(Text-fig. 2). 

It may also be noted in Text-fig. 1 that the populations of the highest altitudes, 
those of the upper levels of Mt. Kenya and of the Kivu volcanos, appear to consist of 
birds a little smaller than might have been expected. If this is genuine it might 
be a result of the following factors : (a) gene-flow, of necessity predominantly from 
lower altitudes, where the birds are smaller ; (b) marginal ecological conditions, 
implying poorer feeding — the hypothesis put forward by Davis (1938) to account for 
the unexpectedly small size of some mountain populations of rodents. 

Wing-length and minimum temperature 

The partial regression equation given above shows that wing-length increases 
as minimum temperature decreases — which is in accord with Bergmann's rule — and 



332 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

that this factor operates more powerfully than the others under consideration. 
Moreover, as shown by the consistent grouping of the points in Text-fig. 2 along an 
approximately straight line, this temperature factor (in conjunction with altitude) 
overrides any ecological adaptation, even though the populations involved occupy 
habitats as different as evergreen forest and dry thorn- bush. The consistency of 
the combined effect is further illustrated by the following facts extracted from 
Appendix 3 : 

(1) Where populations otherwise similar are subjected to different minimum 
temperatures (and altitudes), those birds inhabiting the cooler climate have 
longer wings in 18 out of the 19 groups of comparable populations. (In these 
cases of course the altitude and minimum-temperature factors are reinforcing 
each other.) The only exception occurs in populations 59-64, and there the 
difference on the "wrong" side is extremely small. 

(2) The five savanna populations on the south of the Sahara, nos. 14-18, 
range in wing-average only from 53-5 to 56-1 and the minima from 62 to 67 ° 
F. Yet even within these narrow limits the correlation is so effective that the 
populations with the shortest wings, 53-5 and 537, occur with the highest 
minimum temperatures, 66° and 67 F., and the other three wing-lengths are 
in inverse order of the temperatures. 

(3) On the eastern rim of the Congo Basin, as already noted, on a line some 
70 miles long from Kamituga eastwards (populations 52-54) mean wing-lengths 
fall from 61-4 mm. with minimum 50 (at 7,000 ft.) through 55-9 with 6o° F. 
(at 4,300 ft.) to 53-1 mm. with 66° F. (at 3,000 ft.) — another case in which 
altitude and temperature are operating together. 

The only local peculiarity disclosed by Text-fig. 2 is that the South African 
populations (black circles), grouped as they are to the right, have rather shorter 
wings than would be expected from the low minimum temperatures. Since among 
the African Zosteropidae they alone are outside the tropics, the first suspicion is that 
we have here a " latitude effect " such as Snow (1953) demonstrated in the Paridae. 
He suggested that in the short daylight of winter the high-latitude tits could not 
maintain a body of the size most efficient for heat-conservation. This could hardly 
apply to South African birds, the duration of whose feeding days in winter would 
be only 2-3 hours shorter than those on the equator — and much longer than those 
in the latitude of Britain ; but I can think of no acceptable ecological reason why 
South African Zoster ops should show this peculiarity. 

Among the South African birds two populations are outstanding, namely the 
pallida of the south-west and the (higher-altitude) pallida of the Orange Free State. 
Both have conspicuously shorter wings than might have been expected from the 
low minimum temperatures — which are well based on local meteorological records. 
This effect is the more remarkable because the maximum temperatures are rather 
high and hence favour lengthening of the wing. 

In connection with this peculiarity of pallida two possibilities may be suggested. 
First, the bodies may be stunted as an adaptation to rigorous ecological conditions. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 333 

This seems unlikely because, although the environment of South West Africa, with 
its aridity and extreme daily range in temperature, is one of the harshest experienced 
by Zoster ops, in the Orange Free State, especially along the Vaal River, whence some 
of the specimens come, conditions are not so bad. The alternative possibility is 
that in this Zosterops the wing is abnormally small for the body. No data exist for 
testing this : but these birds show another peculiarity that is presumably not 
independent, namely an extremely high ratio of tail-length to wing-length, as is 
obvious in Text-fig. 5. 

Wing-length and maximum temperature 

The statistical demonstration that the wings of African Zosterops tend to get 
longer as the maximum temperature rises is contrary to Bergmann's rule. It is 
also contrary to the impression given by Text-fig. 3, but this is undoubtedly falla- 
cious and due to the fact that (as shown by the equation) minimum temperatures 
and altitude have, in combination, an effect on wing-length powerful enough 
completely to mask the influence of the maximum temperature. Here, then, in 
Text-fig. 3 we have an outstanding example of the misleading result that can be 
produced by the use of a graphical method unchecked by statistical analysis. 

The tendency for wing-length to increase with a rise in the maximum temperature 
is difficult to explain on biological grounds, because the reverse would be expected. 
It should, however, first be pointed out that temperature is not an adequate index 
of the demands made by the environment on the bird's potentiality for heat exchange, 
because evaporation depends greatly also on the relative humidity of the air. This 
is subject to wide local variation according to the general climate and cannot be 
neglected when the effects of maximum temperatures on heat-exchange are being 
considered. 1 Far too few stations record relative humidities for it to be possible to 
calculate the mean saturation-deficits experienced in the hot season by any of the 
populations under discussion. All that can be said, from a knowledge of the climatic 
regimes, is that more of the very high maxima are accompanied by low humidities, 
and hence are biologically less exacting, than are accompanied by high humidities. 
And it is virtually certain that if the populations could be arranged in order of 
the saturation-deficits they experience in the hot season it would differ a good deal 
from that of their hot-season maximum temperatures. 

In birds exposed to the greatest heat some special adaptations for escaping or 
reducing the full effect may be present — faster breathing with or without gaping, 
a higher ratio of surface (internal and/or external) to mass, thinner plumage, or 
a persistent seeking of the coolest available eco-climates. 

Wing-length and other temperature combinations 

It has been suggested that it might be interesting to ascertain whether statistically 
significant correlations exist between wing-length, annual mean temperature (m) 

1 This does not apply to minimum temperatures, which are practically always accompanied by high 
humidities, if not by dew-fall. 



334 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

and annual range in temperature (r), m being '—^- and t max. — 

min. The partial regression equation with altitude then becomes : 

w = 59-8 -f 0-690 -f- o-2or — o-i6m. 

(Standard errors of both m and r are -04, so that both coefficients are highly signi- 
ficant.) This means that wings tend to be shorter with higher means and to be 
longer with greater annual ranges. The effect of the higher mean would be in confor- 
mity with Bergmann's rule ; and the effect of the higher annual range is what would 
have been expected from the first equation. For annual range is a reflection of 
both higher maxima and lower minima ; and each of these factors was shown to be 
correlated with longer wings. 



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30 32 34 36 38 40 42 44 46 48 50 



TAIL-LENGTH (mm) 

Fig. 4. Wing-length /tail-length correlations. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 335 

TAIL-LENGTH IN RELATION TO WING-LENGTH 
AND ENVIRONMENT 

Mean tail-lengths vary from 31-2 to 48-7 mm. and the correlation between mean 
tail-length and mean wing-length is remarkably high, -f- 0-93 — cf. the grouping 
in Text-fig. 4. Conformably, the equation showing the multiple regression of 
tail-length (t) on altitude, minimum temperature and maximum temperature takes 
exactly the same form as that of wing-length (p. 329), being : 

t — 44-0 -+- 0730: — o-44n + o-2i#. 

Again, these figures are highly significant statistically (standard errors 0-19, 0-05 
and 0-07 respectively) and they mean that the tails tend to increase with altitude, 
with fall in minima and with rise in maxima. These effects can, however, be shown 
to be due almost entirely to the closeness of the correlations between wing-length 

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»•»■•»«• 68 — }5 72 74 76 --78 BO 

TAIL/WING RATIO l0 ° X TAIL 

WING 

Fig. 5. Correlations between wing-length and tail/wing ratio. 



336 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

and tail-length because, when the multiple regression of tail-length is calculated on 
wing-length together with the other factors, the equation becomes : 

t = —30-50 — 0-1200 — *92w -f- 0-062* + 1-246*0. 

Of these the wing factor (w) is highly significant (standard error 0-114), the 
minimum temperature is just significant (standard error 0-45) and the others are 
not significant. 

Reverting to the first of these two equations, and applying the same method 
as was used with the wings (p. 330), it can be inferred that in the African Zoster ops 
the potential total effect of altitude on tail-length is 7 mm., of minimum temperature 
15 mm., and of maximum temperature 7 mm. These figures, 17%, 37% and 17% 
respectively of the overall mean tail-length, are proportionately greater than the 
corresponding effects, 12%, 17% and 7%, of the respective factors on the wing. 
Hence, compared with the wing, the tail is relatively more sensitive to the effect 
of temperature than to that of altitude. This is confirmed by the last equation, 
in which it is shown that minimum temperature has a significant influence (along 
with wing-length) on tail-length, but that altitude has not. This result accords 
with the probability that variations in air-pressure may not exercize a strong and 
direct influence on tail-length ; and certainly not so strong as on wing-length. 

As shown in Text-fig. 5, the tail /wing ratio rises with increasing wing-length ; in 
fact, even omitting the abnormal Cameroons group (marked x in Text-fig. 5), in 
the shortest-winged populations the tails average only about 67% of the length of 
the wings, while in the longest-winged tails average about 76%. x It is shown by the 
last equation that this is unlikely to be due to a direct altitude effect, i.e. to depend 
on air-pressure (cf. the scatter when tail /wing ratios are plotted against altitude 
in Text-fig. 6), but the possibility of a temperature effect is not excluded. Thus, 
the African Zosterops seem to show the same phenomenon as the Palaearctic Paridae, 
in which tails are proportionately longer in colder climates (Snow, 1953). It is 
possible that the same holds good in Malaysian birds generally (Longhurst, 1952), 
but the climatic data cited for these leave something to be desired. This result 
is terminologically at variance with Allen's rule, under which extremities are pro- 
portionately shorter in cooler climates ; but this rule is probably not applicable 
to an appendage such as a bird's tail, in which there is no circulation and the heat- 
exchange is presumably negligible. 

The biological reason for the progressive change in the tail /wing ratio of the 
African Zosterops is difficult to suggest. I owe to Mr. Bernard Stonehouse the idea 
that if, as appears, the Zosterops inhabiting the cooler climates have more ample 
contour feathers, then the other feathers might share in the process of elongation ; 
and owing to the less specialized nature and function of the tail-feathers this process 
might in them be less subject to modification by natural selection than in the 
flight-feathers. 

There are no marked discontinuities in Text-fig. 4 although two geographical 

1 Since the above was written. I have received Mees' paper in Sarawak Mus. J., 6 (1955) : 641-661, in 
which he describes a similar change in tail/wing ratio with wing-length in certain eastern Zosteropidae. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 337 

groups occupy slightly aberrant positions. Moreover, in these two groups the 
alignment of the points is parallel to that of the main body. It can be concluded that 
the same principles are operating in all forms of Zosterops and that there is no over- 
riding adaptation of tail/wing ratio to habitat (in particular, to open thorn-bush as 
against evergreen forest). 



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2000 4O00 6000 9000 IOOOO 

ALTITUDE (FEET) 

Fig. 6. Correlations between tail/wing ratio and altitude. 



338 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

In Text-fig. 4 certain populations, 19-23 and 66 in Appendix 3, lie to the left (open 
circles joined by the broken line), that is, their tails are abnormally short in relation 
to their wings. The segregation of 19-23 is equally marked in Text-figs. 6 and 7, 
where tail /wing ratio is plotted against altitude and minima respectively. All the 
populations concerned lie round the head of the Gulf of Guinea on and near Cameroon 
Mt. (see Map 5), except 66, which is in Tanganyika. Moreover the tail /wing ratio 
of the birds of Fernando Po (x in Text-fig. 4), which are identical in plumage, lie 
on the same line exactly. The biggest bird in this series (wing 63 mm.), from the 



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*0 6S 60 SS SO 45 40 35 

MINIMUM TEMPERATURE (°F) 

Fig. 7. Correlations between tail /wing ratio and minima. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 339 

top of Cameroon Mt., is so generally aberrant as to be placed in a different genus 
(Speirops), but nevertheless its tail /wing ratio is on the same, abnormal, line as 
those of its unrelated neighbours. This suggests that the character " short tail " 
in the other Gulf of Guinea birds may not be of taxonomic significance, but is related 
to some ecological peculiarity of the area. 

The South African populations (usually attributed to at least two different 
species) are not completely separated in Text-fig. 4 (black circles) from the remainder, 
as are those of the Cameroons, but they do all lie to the right, having their tails 
abnormally long for their wings. It seems unlikely that this can be correlated 
with some peculiar environmental factor, because the birds concerned inhabit an 
extremely wide range of habitats (as described in Part 4) . 



BEAK-LENGTH IN RELATION TO WING-LENGTH 
AND ENVIRONMENT 

Correlation between beak-length and wing-length is high, + 0-83. The partial 
regression equation for beak-length (b) on altitude, minimum and maximum 
temperatures is of the same pattern as those for wing and for tail, namely : 

b = 14-11 mm. + 0-1440 — 0*039^ -f- 0-005%. 

In this equation a and n are significant, with standard errors 0-04 and 0-012 
respectively, but x is not. Hence it is inferred that altitude and minimum 
temperature each affect beak-length, in the same way as they do wing-length, but 
to the same potential extent, each up to a total of 1-5 mm. 

If now, as in the case of tails, the partial regression of beak-length is calculated on 
wing-length as well as on the environmental factors, the equation is : 

b = -f 0-44 — 0-0130 + 0-25^ — 0-022% -f 0-2295^. 

Again, the last factor, wing-length, is highly significant (standard error -031), 
the minimum temperature just significant (standard error -012) and the altitude and 
maximum temperature not significant. This means that the over-riding factor in 
determining the length of the beak is the length of the wing (i.e. presumably the 
general size of the bird), but that there is also a slight tendency for beak-length to 
increase with higher night temperatures. This would accord with Allen's rule, 
under which exposed, terminal, parts of the body are proportionately longer in 
warmer climates. 

When beak-length is plotted against wing-length (Text-fig. 8), the data look 
homogeneous. Points relating to birds of dry country are not segregated from 
those of forest. Hence there is no important adaptation of beak-length to type 
of habitat or type of tree. 

In Text-fig. 8 it will be seen that as a whole the southern tropical populations 
(distinguished by a dot in the circle), some of which live in savanna and some in 



340 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

mountain forest, have a beak-length that is nearly constant in spite of differences 
in wing-length. This is interesting because the populations comprising this southern 
group have usually been divided between two species. 

With one exception, the Speirops of the top of Cameroon Mt., the beaks that are 
longest in relation to wings all belong to birds of the highlands of Abyssinia, eastern 
Kenya and north-eastern Tanganyika (squares on Text-fig. 8) , a group of populations 
there are other grounds for thinking closely allied. In this respect (though the dif- 
ferences are very small) they stand apart from the Zosterops of the other high moun- 
tains of Central Africa and western Kenya. For this no ecological reason can be 
suggested, since all the mountains are characterized by evergreen forest and this is 
represented by the same type, though not exclusively, in both areas. 

The two populations with proportionately the longest beaks of all are those 
(otherwise different in both dimensions and plumage) confined to the tops of the 



65 



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00 



BEAK LENGTH (MM) 

Fig. 8. Wing-length /beak correlations. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 341 

widely separated Kulal and Teita Mountains (populations 28 and 34 in Appendix 3). 
Again, no ecological reason can be suggested. The number of individuals on each 
mountain is certainly small, so that in this respect conditions are from the evolu- 
tionary point of view more comparable with that of a bird-population on an oceanic 
islet than of one living in a continent. 



DIMENSIONAL CORRELATIONS AND TAXONOMY 

The one bird of undoubted taxonomic distinctness at a high level, the Speirops, 
is abnormal in only one of these correlations, namely tail /wing ratio, and in that it 
agrees with its immediate neighbours. 

On the other hand, there are four groups that show some peculiarity not explained 
on ecological grounds, which might therefore be regarded as primarily phylogenetic: — 

South African birds (black circles in text-figs.). On the whole these are 
unusually long- winged (? large) for their altitudes and somewhat short- winged (? small) 
for their cold-season minima, especially in the case of pallida. They also tend to have 
high tail /wing ratios and they are segregated from other African Zoster ops when this 
ratio is plotted against altitude (Text-fig. 6) and, less sharply, against minimum 
temperature (Text-fig. 7). In this respect also, pallida is the most divergent of the 
South African birds. 

Southern tropical populations (dotted circles in text-figs.). These birds are 
geographically contiguous with the South African, but they differ markedly from 
them in their tail /wing ratios: the points in Text-fig. 5 are distributed in two parallel 
linear groups. This segregation is noticeable also when tail /wing ratios are plotted 
against altitude (Text-fig. 6) and against minima (Text-fig. 7). Also the southern 
tropical birds tend to have beaks of much the same length while their wing-length 
varies. These facts suggest that the southern tropical birds form a single species 
distinct from the South African birds. 

Cameroons group (crosses in text-figs.). The tail/wing ratio is aberrant (Text- 
fig. 5) not only in itself but also in relation to altitude (Text-fig. 6) and minima 
(Text-fig. 7). But because this peculiarity is shared by the local Speirops it is 
thought not to be of taxonomic significance. 

East and northeast African montane birds. The beak/wing ratio of these 
birds, commonly divided into several species, is consistently high. 

Conclusion. The foregoing analyses disclose correlations of dimensions with 
each other and with environmental factors that are remarkable for their general 
application throughout the continental African Zosteropidae. There are no 
abnormalities so striking as to provide compelling reasons for separating populations 
at the specific level, though peculiarities exist that may serve as ancillary characters, 
to be considered in cases of disputed relationships. 

The dimensions of insular populations, which are given in supplementary tables 
in Appendix 3, do not lend themselves to statistical analysis on the same lines as the 
continental data. The results obtained above give, however, standards of 
comparison, utilized in Part 5. 

ZOOL. 4, 7. 23 



342 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

PART 4 

CONTINENTAL POPULATIONS : 
CHARACTERS AND INTERGRAD ATIONS 

In this part the Zosterops of Africa will be described in six geographical sections 
(see Map 3 for key to local maps), an arrangement adopted in order to assist 




Map 3. Key to area maps. 



exposition of the complicated material. At the end of each section the main out- 
lines will be summarized and the appropriate nomenclature indicated. The taxo- 
nomic arrangement at the specific level is discussed as a whole in the general 
synthesis (Part 6) ; meanwhile, the results have been anticipated in Part 1 and 
sketched in Map 1 . Throughout the sections that follow, the discussion will be in 
the light of the correlations and general principles dealt with in Parts 2 and 3. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 



343 



NORTH-EAST AFRICA 

Throughout Eritrea, Abyssinia and Somaliland the lowlands are dry, with acacias 
and combretums as the dominant trees ; and the east and southeast are particularly 
arid. The highlands still carry patches of evergreen forest, largely in its driest form 
(dominated by Juniperus procera), but richer in the more humid south-west of 
Abyssinia. 

The Zosterops situation is most complicated and interpretation is hindered by the 
lack of field data. Six main forms exist, the habitats and chief characters of which 
are outlined in Table I. The last two (highland) forms have always been regarded 
as specifically distinct, both from each other and from the lowland forms. The 
latter conclusion is no doubt correct, the former probably not (see below). 



Table I. — Main Characters and Habitats of the Zosterops of N.E. Africa. 



Range. 

N. and E., lower altitudes 
(up to 5,000 ft.) 

S. E., lower altitudes 
(up to 6,ooo ft.) 



N.W., lower altitudes 
(up to 6,000 ft.) 

S.E., lower altitudes . 
(up to 6,000 ft.) 

Western highlands 
(4,500-9,000 ft.) 

Highlands elsewhere . 
(5,000-10,500 ft.) . 



Habitat. 

Any arborescent vegeta- 
tion (Erlanger 1907, 
K. D. Smith in litt.) 

Dry acacia country (Er- 
langer 1907). Thorn 
scrub and, exception- 
ally, juniper woods 
(Benson 19466) 

" Steppe " (Neumann 
1906) 

" River valleys " (ibid.) 
Deciduous forest 
(Cheesman in lift.) 

Mountain forest (Neu- 
mann 1906) 

Evergreen forest (cf. 
Benson 19466) 



Name used 
by Sclater 

(1930)- 


Colour of 

A 


r 

Upper 
parts. 


> > 
Belly. 


abyssinica 


Grey-green 


Greyish 


smithi 


Greyish 
yellow-green 


Yellow 


senegalensis 


. Yellow-green 


Yellow 


omoensis 


Yellow-green 


Greyish 


kaffensis 


Rich green 


Yellow 


poliogastra . 


Rich green 


Grey 



The geographical distribution of the six forms is shown diagrammatically in Text- 
fig. 9. It will be seen that a grey-bellied and a yellow-bellied form divide the 
highlands between them, while round them at lower altitudes are arranged four 
other forms, alternately grey-, yellow-, grey- and yellow-bellied. Of the four lowland 
birds the two occupying western Abyssinia, which is damper than the east, are more 
richly pigmented than the others. 

The actual records of the various forms are plotted in Map 4, which gives some 
indication of the complicated relief. Clearly the concept of sympatry and allopatry 
is not easy to apply. Lowland forms penetrate deeply into the highlands: along 
the Rift Valley a line of lowland grey-belly localities separates the eastern from the 
western highland grey-belly populations, while in the south-west the other lowland 



344 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 



X 

x;; 
x - 

x\ 

x; 

X : : 
X 

X'v 
XV 




I'l-jl, 

iiH 1 1 v.zinizi.----- 



K 



- : 'rf++++ + ++ 



LOWLAND YELLOW 
(SENEGALENSIS) 

r++\ LOWLAND YELLOW 
L±ll (JUBAEMSIS) 

II I II HIGHLAND YELLOW (KAFFENSIs) 



+ ^ + + + + 



=-=J LOWLAND GREY (ABYSSINICA) 
LOWLAND GREY (OMQENSis) 



^ 



t— I HIGHLAND GREY (POLIOGASTRA) 
LAND OVER 5000 FT. 



Fig. 9. Distribution (diagrammatic) of Zoster ops belly-colour in N.E. Africa. 



VAR 



IATION IN THE WESTERN ZOSTEROPIDAE (AVES) 345 



X KAFFENSIS 

A POUOGASTRA 

B ABYSSINICA 

9 OMOENSIS 

* SENEGALEfJSIS 

•+• JUBAENSIS 




Map 4. Zosterops distribution in N.E. Africa. 



346 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

grey-belly (omoensis) occupies the Omo River valley, deeply cut through the country 
of the highland yellow-belly {kaffensis). Highland and lowland forms must in 
places actually meet. Two lowlands forms and also poliogastra have been recorded 
from Keren (Reichenow, 1903 ; Zedlitz, 191 1), but this is an area of extremely 
broken country and it may be doubted whether the locality is critically stated. How- 
ever, the specimens (in the British Museum) of both (grey-bellied) lowland omoensis 
and (yellow-bellied) highland kaffensis from Roke suggest overlap there ; and there 
is a specimen of abyssinica from 9,000 ft. (= 2,750 m.), as noted on the label, at 
Bijo, near Harrar, right in the highland {poliogastra) zone. Again, Benson (19466) 
at the extreme south of the range of poliogastra, near Alghe, recorded both this and 
the local lowland birds in juniper woods at 6,000 ft. (= 1,800 m.). One specimen 
from near Keren may be a hybrid between the highland poliogastra and the lowland 
abyssinica (see Note 4, Appendix 1). Finally, Mr. K. D. Smith {in litt.) encountered 
a mixed party of abyssinica and poliogastra at Faghena, on the eastern escarpment 
of Eritrea, on exotic trees in a clearing in the woodlands at 5,570 ft. This is an 
abnormally low altitude for poliogastra and the whole locality is peculiar ecologically. 

The highland birds 

In the highlands the grey-bellies {poliogastra) have a more extensive range than 
the yellow-bellies {kaffensis ; see Note 5, Appendix 1). They occur all down the 
spine of the Eritrean and Abyssinian highlands, west over the plateau to near Lake 
Tana, east along the mountains towards Harrar and south to the end of the high 
country, at Yavello, east of the tip of Lake Rudolf. Grey-bellies everywhere occur 
on both sides of the line of the Abyssinian lakes, but they do not cross the deeply- 
cut valley of the Omo River just to the west. Beyond this the high-level Zoster ops 
are yellow-bellies, which range from Lake Tana southwards. 

The two forms apparently overlap a little. Both occur at Addis Ababa, with the 
yellows much the commoner (Guichard, 1950, and in litt.), and in the neighbouring 
Menengasha forest (B.M. specimens). The Addis overlap may be a recent re- 
colonization following tree-planting of the formerly deforested area. Away to the 
north-west, south of Lake Tana, the Dembacha grey-belly (B.M. specimen ; breeding 
condition), which is partly surrounded by yellow-belly localities, may indicate another 
limited area of overlap. 

Both the grey-bellies and the yellow-bellies show some geographical variation. 
The former are a trifle smaller in the extreme southwest of their range (mean wing 
61-5 mm. compared with 63-9). For an alleged plumage-difference between northern 
and southern grey-bellies, see Note 6, Appendix 1. Of the highland yellow-bellies 
(see Note 5, Appendix 1), those south of Nono are the most richly coloured, both 
above and below, and those from Shoa (the neighbourhood of Addis Ababa) which 
have been distinguished subspecincaHy, are the biggest, and a little duller generally, 
without such large and bright yellow foreheads. But, again, those from Nono and 
further north, within the bend of the Big Abbai (Blue Nile) are intermediate in these 
characters. Thus the variation is not a simple geographical cline and moreover 
plumage-colour is not correlated with rainfall. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 347 

I believe that the grey-bellies and the yellow-bellies of the highlands are con- 
specific. In the middle of their range both forms are the same size, with the same 
tail /wing ratio, and are similar in plumage (including eye-ring), except for the belly. 
Pending further field data, it seems that they form a parallel to the cases of the grey- 
bellies and green-bellies in the Cape (see section on " South Africa " below). True, 
in Abyssinia there is no evidence of interbreeding, but the ornithology of the 
country is such that it could hardly be expected at the present stage. 

With their sharply defined yellow foreheads and comparatively long tails (ratio 
75) the yellow-bellies of the western Abyssinian highlands differ from the nearest 
montane birds, on the Imatong group on the Sudan border, which have dimmer 
foreheads and shorter tails (70). The Abyssinian birds are more like some Kenya 
highlands birds, and though they differ from the nearest of these in having stronger 
carotenoid these appear taxonomically closer than the Imatong birds. 

The lowland birds 

As already mentioned, there are four main forms arranged round the highlands, 
grey-bellies and yellow-bellies alternately. In these birds also belly-colour has 
always been taken as a specific character but, as shown in Part 2, there is no reason 
to think this is correct and their true relationships are difficult to decide. All four 
forms are, as expected, smaller than the highland birds (see Appendix 3). The two 
eastern ones are particularly dingy in plumage, with reduced carotenoid and a 
tendency to muddy brownish wash on the underparts. These characters are not 
shared by the two western birds, the yellow-bellied senegalensis , which reaches Lake 
Tana from West Africa, and the lowland bird of south-west Abyssinia, with its clear 
grey belly (omoensis) . 

The difference in general colour tone between the eastern pair and the western 
pair of birds is connected with the fact that, apart from stronger carotenoid in the 
west, the melanin in these birds is of black (A) type, while in the eastern pair of birds 
it is of the browner (B) type. As discussed in Part 2, the taxonomic value of this 
character is uncertain. In so far as it depends on the humidity or aridity of the 
environment, as in South Africa, it is worthless. But in the West African sene- 
galensis it has been shown that the melanin is of (A) type even in the driest environ- 
ment. The occurrence of the same character in the contiguous omoensis might 
therefore mean taxonomic affinity ; but since omoensis inhabits the wettest environ- 
ment of all the lowland birds, it might alternatively be, as hitherto believed, a form 
of abyssinica, climatically modified. 

Geographical relations between lowland forms 

The grey-bellies of the eastern lowlands extend from the Red Sea Hills (Erkowit) 
of the Sudan " wherever there is thick vegetation " (Cave and Macdonald, 1955) 
all down the eastern foot of the Eritrean-Abyssinian plateau ; and thence through 
the Rift Valley nearly to Lake Abaya and east through British Somaliland. They 
also occur down the western side of the Eritrean and Abyssinian highlands as far 



3.|« VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

as Lake Tana and the beginning of the Blue Nile (Great Abbai) valley. Here they 
seem to overlap the yellow-bellies (senegalensis) from the west, which find their 
eastern limits on the western slopes of these Eritrean and Abyssinian highlands. 
Both forms have been collected near Keren and also on the shore of Lake Tana (see 
Note 7, Appendix i), while the yellow-belly has also been taken on the slopes of 
Mt. Geech, where on geographical grounds the grey-bellied poliogastra (or even 
abyssinica) might have been expected, as will be seen from Map 4. 

The other, south-western, grey-bellied form, omoensis, is typically represented 
from the northernmost point of the Omo River (Nono) southwards through the 
comparatively humid valleys in Jimma and Kaffa to the beginning of the natter 
ground at the head of Lake Rudolf. Specimens from north of Nono are more 
puzzling, Gomit River birds (about fifty miles S.W. of Lake Tana — R. E. Cheesman, 
in litt.) are more like abyssinica, yet a bird from further north, on the west shore of 
the lake (Azobahr) is almost typical omoensis. Two more birds from Kunkur, 
still further to the north and west, are also more like omoensis than abyssinica, but 
they have dull upper parts — in other words, they are in this respect intermediate 
between the two forms, and these birds might be hybrids. But again, the dullness 
in plumage might be correlated with the fact that their environment is drier 
than normal for omoensis (see Note 8, Appendix 1). However, against both 
these hypotheses, Dr. Auber finds that the melanization of these dull-looking 
Kunkur birds is the typical (A) of omoensis without infusion of (B). In any case, 
although omoensis and abyssinica have not been recorded from the same locality 
anywhere, Map 4 suggests that round the west of Lake Tana they may overlap 
considerably ; the big gap in the recorded distribution of grey-bellies just south of 
the Lake might be due to the accidents of collecting — much of the Blue Nile gorge is 
inaccessible. 

On the whole, the ranges of grey-bellied abyssinica and omoensis and yellow- 
bellied senegalensis do not help to settle their taxonomic relationships. The extent 
of their geographical overlap and the absence of indubitable hybrids would in most 
groups of birds suggest specific separation, but in Zosterops the South African situa- 
tion, described subsequently, shows that the ordinary rules do not necessarily apply. 

On the east side of the mountains the widely-ranging lowland grey-bellies, abys- 
sinica, are, so far as known, strictly allopatric with omoensis. This is not recorded 
east of the Omo Valley, while abyssinica is not known west of the Rift. A narrow 
ridge of highland, occupied by the montane poliogastra, intervenes between them. 
Also on the south of its range omoensis appears to be allopatric with the yellow- 
bellies (jubaensis ; formerly smithi) occupying the arid country from the head of 
Lake Rudolf eastwards. Further, at least in Ethiopia these yellow-bellies seem 
allopatric with abyssinica on the north, though from lack of collecting it is not 
certain whether this relationship is maintained eastwards all the way through 
Somaliland (see Note 9, Appendix 1). 

To summarize : in the east and south of Ethiopia abyssinica, jubaensis and 
omoensis are all allopatric ; in the west abyssinica, omoensis and senegalensis all 
overlap round Lake Tana. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 349 

Geographical variations within the lowland forms 

The local representatives of senegalensis (sensu stricto) are typical in colour, but 
the few specimens available from the Lake Tana-Mt. Geech area are abnormally 
large, as expected from the altitude. Their wings measure 59, 59 and 60 compared 
with 52-59 (but mean only 54*6) in the nearest (Sudan) birds at an average of 2,000 
ft. lower. 

The variation in the southern yellow-belly, jubaensis (see also Bull. Brit. Orn. 
Club, 72 : 50-51), is slight and in accord with expectation. Birds from higher 
altitudes, near Yavello, are a little bigger (wing 53-6) than they are further west, 
round Lake Rudolf (wing 52-1), or further east towards the Indian Ocean (wing 51*8), 
and perhaps not quite so dull-plumaged. 

In the more widely ranging abyssinica the variations are more marked. In the 
first place, the most northerly birds on the African mainland, from Eritrea and the 
Red Sea Hills of the Sudan at an average altitude of 3,500 ft., are the smallest, wing 
54-4 mm., compared with Abyssinian birds, ca. 4,500 ft., average 58-3 and British 
Somaliland, ca. 4,000 ft., average 57-1. All specimens from Abyssinia, Eritrea 
and the Sudan have brown beaks, a character unknown elsewhere in Africa except 
in omoensis. But all the British Somaliland birds have beaks more or less black. 
In plumage the northernmost birds tend to be more yellow-green above, not so dull 
(greyish) as the others. 

Two populations of these grey-bellied birds are isolated by water, but both are 
little differentiated. Across the Red Sea birds of this type occur between about 
1,500 and 5,000 ft. from near Mecca (that is, further north than the most northerly 
occurrence of Zosterops in Africa) to the Aden Protectorate, and they have been 
separated under the name arabs. In these birds the carotenoid is further reduced, 
so that they are even more dingy than Abyssinian abyssinica, with no yellow on 
the head and only slightly whitish lores. Moreover, they share the peculiarity of 
the brown beak. In dimensions they are extremely close to both Abyssinian and 
British Somaliland birds — wings 57-5 compared with 58-3 and 57-1 respectively, 
tail/wing ratio 73-5 compared with 72 and 73-4. 

On the " continental " island of Sokotra, off Cape Guardafui, the tip of Somaliland, 
the birds average slightly brighter (more yellow-green) above than continental birds, 
though the yellow on the throat is no stronger, and they have little or no yellowish 
on the forehead. With wings averaging 56-1 and tail /wing ratio 71-8 they are not 
distinctive in dimensions. The beak is scarcely longer in proportion than in the 
mainland populations (beak /wing ratio 24-1 compared with 22-9-23-7), but it is 
interesting that it is not brown but black, like that of its nearest neighbours, in 
Somaliland. Thus the two isolated populations of abyssinica, though named, are 
" poor races ". 

Provisional taxonomic conclusions 

(1) The yellow-bellied and the grey-bellied highland birds, kaffensis and polio- 
gastra are conspecific. It is not worth while to retain the name schoana as distinct 
from kaffensis. 



350 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

(2) The grey-bellied lowland abyssinica is conspecific with the yellow-bellied 
jubaensis (smithi) and the grey-bellied omoensis. 

(3) The yellow-bellied bird that enters western Abyssinia from West Africa 
(senegalensis) presumably belongs to a species different from (1), which looks so 
different, and from (2), which it overlaps. 

(4) The name socotrana is retained, on the character of the black, instead of brown, 
beak, but the range of the form is extended to include British Somaliland. 

REMAINDER OF NORTHERN TROPICAL AFRICA 

The whole northern savanna belt of Africa, south of about 14 N., is occupied 
by a brightly coloured Zosterops (senegalensis) with much yellow pigment and less 
melanin than any other form in Africa. It extends from the Atlantic (Senegal) 
almost to the coast of the Red Sea, from which its range is separated by the highlands 
of Eritrea and Abyssinia (Maps 1 and 4) . Southwards the brightly coloured birds 
extend for an average depth of some five hundred miles towards the Equator with 
very little variation. Almost the whole of this vast area of nearly two million 
square miles is lowland, below 3,000 ft. The main exceptions are (a) the chain of 
small detached areas of plateau and mountain at the head of the Gulf of Guinea, 
(b) the Imatong group of mountains on the Sudan-Uganda border, (c) the western 
slopes of the Abyssinian highlands. (There are also the Futa Jalon massif in the 
west and the Bauchi plateau in eastern Nigeria, from neither of which is any 
Zosterops specimen available.) 

The climate is driest in the north — predominantly thorn-bush country, merging 
southwards into " savanna " (tall grass with small deciduous trees) and ultimately, 
in the wettest parts, into tropical evergreen forest. The highlands carry some 
montane forest. In many parts of the area Zosterops seem to be uncommon and 
only about 200 specimens have been available for study. 

From east to west through the belts of thorn country and savanna no local 
differences in plumage can be detected, though there are interesting differences in 
size (see Appendix 3, populations 13-18). Where, however, the rainfall is higher 
there are changes in plumage, chiefly connected with increases in melanin. Of the 
typically yellow senegalensis the most westerly representatives, from the Gold Coast 
to Senegal (Map 5), are the smallest — wing mean 53-5 mm. In Nigeria, the 
Cameroons and Oubangi-Chari the birds north of about 8° N. are larger, averaging 
55-9, but in Southern Nigeria they are not (wings only 537). Further east, in the 
Sudan, birds north of 8° N. are about the same size (56-1) as those in the same 
latitudes in Nigeria and the Cameroons, but nearer the Equator, in the southern 
Sudan, extreme north-eastern Belgian Congo and the neighbouring corner of Uganda, 
they are smaller (54*6), much as in same latitudes in West Africa. As already noted 
in Part 3, these differences are closely correlated with the minimum temperatures. 
Then from slightly higher altitudes in the flat country of northern Uganda the few 
specimens available are a little larger than the foregoing, and finally the eight high- 
altitude specimens from 5,000-7,000 ft. on the western edge of the Abyssinian plateau 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 



351 






* 

S 
- 1 

> \ 




\ 






\ 




K 


/ 








1 


3H 


\ 


a 


% 


1 


* ' 




1 






1 
\ 


"^ 





352 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 




E T H I O PI A 



,-MT KULAL 



J/IARSABIT 



•Nairobi 



■&NQ\DO MT \£HYULU MTS. 
.•h-KETUMBEIN^,,. 
S^.-MEROt- KJDMANJARO 



ESSIMINGOR MT \_ CtETTA MTS. f 


.-'•"■:. 


:. NORTH ( 


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fyC?^ UFIOME "' 


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..;..; rlANANp K 


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n 



Contour at 4000 ft. 
Map 6. Kenya and Northern Tanganyika. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 353 

and on the slopes of the Imatong group (Sudan-Uganda border; Map 6) are, as 
expected, the largest of all, with wings averaging 59-6 mm. 

Apart from birds so much darker that they have received different names (dealt 
with below), the specimens can be divided into (a) the yellowest, (b) the greenest, 
(c) intermediates — the whole colour-range being small. The yellowest specimens 
are scattered through the whole range of senegalensis. The greenest birds, which, 
along with a few of the intermediates, can be matched with exceptionally bright 
specimens from the savanna belt south of the Equator (anderssoni) , come from the 
high eastern localities just mentioned as providing the biggest birds. 

Within West Africa the Zosterops change more markedly in plumage in the south- 
west corner, where the rainfall rises rapidly to over 120 inches. From the wet, 
forest, areas (rainfall over 80 inches, well-distributed) of Sierra Leone, Liberia and 
Cote dTvoire (Abidjan, rainfall yy inches) all the specimens available are dark and 
dull, apparently owing to a general increase in melanin — i.e. as expected under 
Gloger's rule — without strengthening of carotenoid. (These birds have usually 
been called leoninus of Sclater, but as Rand (1951) has pointed out, this is a synonym 
of demeryi of Biittikofer.) These dark birds have wings even shorter (52-8 mm.) 
than the brighter birds to the north (53-5) and again the minimum temperature 
(68° F.) is exceptionally high. No detailed evidence is available of transition from 
the typically yellow senegalensis to the greener demeryi, though one specimen 
(" Kamasigi, Sierra Leone ") is intermediate. This lack of evidence is probably 
because collecting has been sporadic in this part of Africa and the belt of transition 
from dry country to wet is narrow. 

East of the western Cote dTvoire, drier country comes right to the coast 
again until Nigeria is reached. Here dark birds can again be expected, but the 
present evidence from Nigeria is inconclusive (see Note 10, Appendix 1). Further 
to the south-east, however, round the Gulf of Guinea, the lowlands, which are excep- 
tionally wet and hot, are occupied by the birds of the expected type (the so-called 
pusilla), small and richly coloured, with plenty of melanin giving them green upper 
parts and flanks, but also strong, slightly orange-tinged, carotenoid and a golden 
forehead. With wings 49-53 (mean 51-3), tails 30-33 (mean 31-5), these birds are 
the smallest in Africa — in accord with the high local minimum temperature, 69 F — 
and they also have the lowest tail /wing ratio (61) except for the similar neighbouring 
birds from Manenguba (see below). Birds of this type occupy the southern 
Cameroons and Gaboon south to at least i° 45' S. (Mbigou) and east to at least 16 E. 
Most of this area receives between 70 and 120 inches of rain, well distributed, and 
evergreen forest abounds. It is, in fact, on the north-western edge of the great 
Congo forest ; and similar birds probably exist in a narrow belt along the edge of 
the forest right across the north of the Congo Basin. However, the only skins from 
this belt, which come from Medje, a thousand miles to the east, though recorded as 
pusilla (Chapin, 1954), differ somewhat — see below — so that if specimens become 
available from intervening localities they may be expected to show an east-west 
cline. 

At their western end these little "pusilla" abut upon the mountainous area 



354 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

along and close to the Nigeria-Cameroons border, with Cameroon Mt. (13,000 ft.) 
on the shore of the Gulf of Guinea and 100 miles to the north-east the Bamenda- 
Banso highlands (7,000 ft.), which tail off northwards past Genderu into the flats 
round Lake Chad. Between Cameroon Mt. and Bamenda smaller isolated highlands 
occur — Manenguba, Kupe, and the Rumpis. Serle (1950, 1954) has shown that all 
these highlands are occupied by richly coloured Zosterops (with plumage like that 
of the lowland " pusilla "), which have all been called stenocricota, a form described 
from Cameroon Mt. Also in these mountains several Zosterops have been collected 
that are so dull and dark that they have been regarded as a different species (Z. 
phyllica). They appear, however, to be merely cases of individual deficiency in 
carotenoid (Moreau, 1953). 

The specimens from the highest average altitude, 6,000 ft. in the Bamenda-Banso 
highlands, are the biggest — wing-mean 58-7 mm., tail/wing ratio 65*5. From an 
average of 4,000 ft. on the Kupe, Manenguba and Rumpi Mts. the birds are much 
smaller, wing-mean 53-1, tail /wing ratio 58. The birds of Cameroon Mt., which 
occupy it up to over 9,000 ft., may well be larger at the higher levels, but those 
specimens labelled with an altitude come from an average height of only 4,000 ft. 
and average small — wing-mean 54-2 mm., tail /wing ratio 62. Each of these popula- 
tions has a tail disproportionately short when compared with birds of similar wing- 
length anywhere else in Africa, as already discussed in Part 3. The biological 
significance of this peculiarity, localized round the head of the Gulf of Guinea, 
is unknown. 

While the strong melanin of these Cameroon birds, both montane and lowland, is 
regarded as correlated with the high rainfall, the general appearance of their plumage 
differs from that of the other dark, high-rainfall, Zosterops of West Africa, those of 
Sierra Leone and Liberia, in being brighter, evidently as a result of stronger carotenoid 
throughout. The highland birds are indeed so like some on mountains in East Africa 
that some workers, though not Bates (1930) nor Serle (1950), have regarded them as 
conspecific and not closely allied to the surrounding senegalensis. This conclusion 
would fit with the remarkable affinity between the montane avifauna of Cameroon 
Mt. as a whole with that of East Africa. Half the passerines of Cameroon Mt. belong 
unquestionably to East African species and, though no geographically intervening 
specimens are known, some of the birds are not even subspecifically distinguishable 
— see Moreau (1952) for discussion. 

The idea of specially close affinity between stenocricota and montane birds else- 
where must, however, be rejected. In the first place, the Zosterops (stenocricota) of 
the mountains differ only in size — and that, no doubt, clinally — from the Zosterops 
("pusilla') of the lowland forest at their feet. Actual transition to typical yellow 
senegalensis is shown so far only by three specimens which are intermediate in both 
colour and tail /wing ratio between stenocricota and senegalensis ("genderuensis" 
Reichenow), from 100 miles north of the Bamenda highlands, where the country 
is somewhat lower and drier. Collecting in the lowlands east and west of Bamenda 
may be expected to yield further intermediate specimens; but owing to the closeness 
of the isohyets the belt of transition is probably narrow. To the west no specimens 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 355 

are at present known from nearer than Enugu (100 miles away), a savanna locality 
where the birds are typically yellow senegalensis , and to the east the situation is 
rather similar (see Note n, Appendix 1). 

Cameroon Mt. differs from every other in the African continent in possessing two 
Zosteropids of undoubtedly different species, for its higher parts are inhabited by 
both stenocricota and the highly aberrant melanocephala, which appear fully sympatric 
and which associate together on the trees (Serle, 1950). With its black and brown 
plumage, pale beak and general loss of carotenoid, melanocephala differs so greatly 
from all others on the African continent that it has usually been kept in a separate 
genus, Speirops, along with other aberrant birds on the islands in the neighbouring 
Gulf of Guinea. There also each Speirops occurs alongside a more normal Zosterops. 
These are discussed in Part 5 below, and because the situation on Cameroon is so 
like that on the islands discussion of melanocephala will be postponed. 

Further east, through the southern zone of the savanna and as far as the north- 
eastern Belgian Congo and Uganda, typical "yellow" senegalensis persist without 
change of plumage to south of 3 N. There incipient change can be detected. In 
the Congo (Map 7), while birds from Niangara (3 35' N., 27 50' E. ; in savanna) 
appear typical, those of Garamba (4 10' N., 30 E.) and also Ekibondo (3 32'N., 
28 24' E.) are a trifle greener, while at Medje (2 26' N., 27 17' E.) in a wetter, 
forest, climate, the birds are much darker. As already noted, they have gone by 
the same name (pusilla) as the equally small birds a thousand miles to the west, 
but at Medje they are not quite so short-tailed (ratio 65 against 62) and are much 
less yellow on the forehead. Zosterops do not seem to extend further south towards 
the centre of the Congo Basin (cf. the blank shown on Map 1). Chapin (1954) living 
for a year at Avakubi, eighty miles nearer to the equator, never recorded a Zosterops. 
As already noted, in the extreme east of the range of senegalensis, on the borders 
of the Sudan, the birds become greener as they ascend the mountains on the Sudan 
border (the Imatongs, Didingas and Dongatona). The Zosterops associated with the 
forests on the tops of these mountains have nearly as much melanin as those on 
the mountains of the Cameroons, though not quite such rich yellow, and are in- 
distinguishable from birds on the mountains round Lake Nyasa, over a thousand 
miles to the south and on the other side of the equator. This occurrence of "yellow" 
and "green" Zosterops on the same small mountain will be further considered when 
the more numerous examples in the Nyasa area come to be dealt with. Meanwhile, 
it seems that the green Imatong birds, like those of the Cameroons highlands, are 
derived from the surrounding "yellow" senegalensis and owe their resemblance to 
each other and to the Nyasa birds to convergent evolution in similar climates. 

South of the Imatongs, in Uganda north of about i° N. (which is dry), the 
Zosterops are a trifle greener than typical senegalensis. It seems certain that this 
indicates transition to the distinctly greener birds, long known as stuhlmanni, in 
the country with higher and better-distributed rainfall on the northern and western 
shores of Lake Victoria (Maps 6 and 7) . These birds differ in plumage from those of 
the tops of the Imatongs (and elsewhere) only in a slightly warm tinge, which has 
been described as "cinnamon". More collecting is needed to establish this transition 



356 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 




Sparse dotting indicates 
s country above 1500 m. 
(4900 ft.), close dotting 
above 2000 m. (6600 
ft. 



Map 7. Central Africa. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 357 

in Uganda thoroughly (see Note 12, Appendix 1) and some discontinuity may be 
expected from the long east-and-west barrier interposed by the swamp belt of Lake 
Kioga and associated waterways. This transition is important in interpreting the 
taxonomy of the African Zosterops, because, as will be shown below, the Lake 
Victoria birds lead gradually to populations with plumages as richly green as any 
in Africa. 

Summary 

Bright yellow Zosterops extend from Senegal to the western slope of the Abyssinian 
plateau. In size they vary consistently with climate and altitude, always averaging 
larger in higher and/or cooler localities. Wherever the rainfall in the lowlands is 
such as to give a "forest climate", the birds show more melanin ; but the resultant 
plumages are slightly different in the four detached wet areas, Sierra Leone — Liberia, 
head of the Gulf of Guinea, north-eastern Belgian Congo, and southern Uganda. 
Transition from "yellow" to "green" birds appears to take place in all these cases 
in a few miles distance, in conformity with rapid change in climate. The two mountain 
groups that are surrounded by "yellow" senegalensis , at the head of the Gulf of 
Guinea and on the Sudan-Uganda border, are both occupied by larger and greener 
birds. Here again, transition from the lowland, yellower, type is believed to take 
place. 

It is concluded that (except the aberrant bird of the top of Cameroon Mt.) the 
whole of the birds considered in this section belong to the same species, senegalensis ; 
the subspecific names that can usefully be applied are demeryi for the greener birds 
of Sierra Leone, stenocricota for the greener birds round the head of the Gulf of 
Guinea and eastwards and stuhlmanni for the greener birds on Lake Victoria. 

KENYA AND NORTHERN TANGANYIKA TERRITORY 

This area (Map 6) differs from all others considered in the number and abrupt 
isolation of its ecological (montane) islands and, conformably, in the number of its 
well-marked forms of Zosterops (Text-fig. 10) within a small area. The lowland birds 
have always been regarded as specifically distinct from the highland, which is 
doubtless correct, and there has been great diversity of opinion about the specific 
allocation of the highland birds. Essentially, the area consists of dry thorn-bush 
and "savanna", nearly all with rainfall under 30 inches, out of which rise the Kenya 
Highlands and an archipelago of smaller highlands, each bearing some montane 
forest, but effectively isolated from each other by lower, hotter and drier country. 

The Kenya Highlands extend about 150 miles each way, at altitudes of at least 
5,000 ft., with several parts rising to 10,000 ft. and over. Much montane forest, 
which has certainly been more continuous in recent historical times, still exists 
throughout the Kenya Highlands, but they are deeply penetrated by savanna. In 
particular the Rift Valley past Nakuru and Naivasha is far drier than the moun- 
tains on either side ; and although it is narrow and much of its floor is over 5,000 ft. 
a.s.L, it is something of a barrier for forest organisms. 

ZOOL. 4, 7. 24 



358 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES; 



too miles 






#B/ KULALENSIS 




„ .. #•□/ 




•P/l) / 

\ / 


''"': 


\ / 


JACKSON/'^O/ 

? 

? 
? 


V/ 

J. j JACKSON t 

, • * • < /" 


••d/ ,oD/// 




•DA 




' -A OQ/// K t K UYUENSlS 




'■■■ /'/"; ■••' 




>D/j 


O Yellow-green belly 




B Grey belly 




OV Breadth of 


6 D/ /"\....,_« / 


®) eye ring 

'1 

//> Extent of yellow 


\'©D ^c^' C / '-'s ®MS/LVANA 
/_... •-. MBULUENS/S 


III) on forehedd 






O D // O ■ / JV/ W/ FREDAE 


1 Connects populations 


MBULUENSIS ' : :'•. 


j o f sim ilar plumage 


•a/V..;. 


1 that are separated 


STIERLINGI 1 
f 


J by ecologically unsuitabk 




1 country 





Fig. io. Geographical distribution of characters in East African Zoster ops. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 359 

The less extensive highlands of East Africa range in size from Kilimanjaro, which 
rises to nearly 20,000 ft. and carries a great girdle of forest, to Kasigau and Marsabit, 
each a little over 5,000 ft. with a forest cap, and the anomalous East Usambara 
plateau, with a forest covering that, thanks to proximity to the Indian Ocean, 
extends practically down to sea-level. 

The dry country separating the various highlands is occupied by a small, rather 
pale, yellow-bellied Zoster ops that surrounds all the montane forms. This type of 
bird is palest and greyest in the north-east, where, on the Abyssinian border, we have 
already met with it, from Lake Rudolf to the Indian Ocean, under the name jubaensis. 
Further south, down the coast 1 and also inland, birds of this type become slightly 
greener above (less grey) as the altitude and rainfall increase. Finally, in the neigh- 
bourhood of Nairobi and again in Mbulu,where they reach as high as 6,000 ft. a.s.l., 
these non-forest Zosterops (here usually called flavilateralis) are birds of yellow-green 
upper parts and lemon-yellow underparts with some green on the flanks. Even at 
their brightest and freshest they have, however, a certain "dusty" look, shared by 
no other yellow-bellied Zosterops on the continent. As noted previously, this effect 
is probably due to the nature of their melanization (B type). 

These dry-country birds increase in size with increasing altitude, as expected. 
The specimens from below 4,000 ft. (average altitude 2,500 ft.) have mean wing- 
length 54-0 mm., those from higher altitudes (averaging 5,000 ft.), 55-3 mm. It 
may be added that in this dry habitat the Zosterops probably move about a good 
deal, with corresponding increase in gene-flow. 

These savanna Zosterops and the montane birds that they surround can hardly 
be regarded as geographically separated, for on the slopes of the mountains savanna 
and secondary bush interdigitate with the forest, which, owing to human inter- 
ference, is nearly everywhere in retreat. Moreover the ecological separation between 
montane forest and savanna Zosterops is not complete, since both are liable to 
frequent secondary growth and introduced trees. Mr. A. Forbes- Watson in litt. has 
noted that in the Teita Hills at over 5,000 ft. both the local mountain birds (silvana) 
and the savanna birds frequent the same (non-primary) vegetation and that the 
same species of fruit can be found in the stomachs of both. Actual mixed feeding- 
parties of highland and lowland Zosterops have been noted in a garden near Nairobi 
(kikuyuensis and flavilateralis, Mr. J. G. Williams in litt.) and in exotic trees in Mbulu 
(nibuluensis and flavilateralis in Grevillea used as coffee-shade, Dr. G. Zink in litt.). 

There is no suggestion of hybridization between the two forms and no difficulty 
in distinguishing the montane birds from those of the surrounding savanna anywhere 
in Kenya or Northern Tanganyika except in Usambara. There montane and savanna 
birds begin to converge in appearance, with a consequent problem of differentiation 
that continues southwards through Tanganyika Territory. 

So far as known, all the East African mountains that retain any forest are in- 
habited by montane Zosterops (see Note 14, Appendix 1), even such small and 

1 It appears to be a fact, and one for which no explanation can be offered, that in southern Kenya 
and all through Tanganyika Territory, Zosterops are extremely rare in the comparatively damp coastal 
strip, the vegetation of which appears to be thoroughly suitable for them (see Note 13, Appendix 1). 



360 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

isolated mountains as Marsabit and Kasigau. Their differentiation is remarkable 
for, apart from smaller variations, there are eight forms so unlike in appearance 
that they have been allocated among five different species. Five of the eight forms 
occur east of Kilimanjaro, within a space of 250 miles ; and of these five each is 
within sight of the stations of at least two others. Two of the geographical gaps do 
not exceed twenty miles. Thus, although the differences between the forms are not 
so great, the Zosterops situation in this ecological archipelago of East Africa is 
comparable to that of the rendovae group, isolated from each other by sea, in the 
Solomon islands, to which Mayr (1947) has drawn attention. 

In the East African montane birds the main variation is in three characters, the 
distribution of which is shown diagrammatically in Text-fig. 10 : 

(a) The birds of Kulal in the north, Teita and South Pare (N.E. and S.E. 
respectively of Kilimanjaro) have grey bellies. All the rest have bellies more or 
less green and yellow. 

(b) Eye-rings are biggest in Kilimanjaro and Teita birds, up to 5 mm. broad 
and covering much of the side of the head ; smaller but still bigger than else- 
where in Africa (up to 3 mm. broad) in the eastern Kenya highlands and in an 
arc round the north of Kilimanjaro, Mbulu-Chyulu-North Pare ; smallest in 
the north, northwest and southeast. 

(c) Golden foreheads are particularly large and rich in the eastern Kenya 
highlands ; somewhat less marked in Mbulu-Chyulu-North Pare ; entirely 
absent on Kilimanjaro and Teita ; and intermediate elsewhere. 

In Kenya west of the Rift the highlands (above about 6,000 ft.), from Mt. Elgon 
to Loliondo, are occupied by a large, rather dull-coloured green bird (jacksoni) with 
rather small eye-ring and a yellow forehead that is usually narrow but sharply 
defined. The carotenoid throughout the plumage is rather "colder", less golden, 
than that of the other montane Zosterops of East Africa. The most northerly birds, 
those on Elgon, tend to have less sharply-defined foreheads (so that some females 
look like South African and Nyasaland birds) and to be slightly yellower than those 
of the main West Kenya Highlands. (But I agree with those who hold that they do 
not merit the separate name elgonensis.) These tendencies, if slightly exaggerated, 
would produce a bird like those inhabiting the tops of the Imatong group of moun- 
tains on the Sudan border (top left-hand corner of Map 6), about 200 miles to the 
north of Mr. Elgon. This Imatong type is believed to have arisen there by local 
evolution (see preceding section) and the resemblance is to be borne in mind when 
considering the classification of the African Zosterops as a whole. 

Within the western Kenya highlands the size of the birds varies, at least in part 
as expected from the altitudes. The largest birds (wing 62-0 mm.) are in the centre, 
between Elgon and Naivasha and including the Mau, at a mean altitude of 8,500 ft. 
Further south, mean altitude 7,000 ft., wings average only 60 -6 mm. Also in the 
north, from Mt. Elgon itself, the birds are small (wing 6o-6), but the few specimens 
available with altitude data average only 7,000 ft. 

It is not certain what relationship the jacksoni of the western Kenya Highlands 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 361 

bear to the much yellower stuhlmanni of the shores of Lake Victoria, a bird regarded 
in the preceding section as transitional from the yellow senegalensis of West Africa. 
From Mr. Elgon westwards for at least 100 miles no Zosterops seems to be known in 
collections. However, south of Elgon, from an area less than 100 miles square, 
mostly at about 4,500-5,000 ft., round Kakamega and the basin of the Yala River, 
the birds (described by Van Someren as yalensis) are brighter than jacksoni, with 
less melanin generally, and are smaller (wing 58-3 compared with 62-0). In their 
slightly yellower plumage and, in some individuals, less defined yellow on the fore- 
head, the Yala birds show a tendency towards the Lake Victoria birds. No cline is, 
however, demonstrable because nearly 150 miles intervene between the Yala 
localities (see Note 15, Appendix 1) and the nearest stuhlmanni specimens. 

Birds of jacksoni type extend east across the Rift Valley in a puzzling fashion, 
which does not seem to have been realized hitherto. This will be reverted to later. 
Meanwhile the point to make is that most of the Kenya Highlands east of the Rift 
are occupied by a strikingly different bird, kikuyuensis, one of the most showy 
Zosterops in Africa, with very large golden patch on the forehead and a broad white 
eye-ring. Apart from the usual larger size at higher levels, the Mt. Kenya birds are 
indistinguishable from those further south, to beyond Nairobi, and from the 
Aberdares (see Note 16, Appendix 1). It is unfortunate that the various specimens 
in collections labelled "Aberdares" bear no precise localities or altitudes, because 
somewhere towards the north end of the range there seems to be contact with birds 
of jacksoni type, and it is important to know what their relations are. 

Typical jacksoni come right to the wooded western edge of the Rift Valley, in- 
cluding the Eburru spur, which projects just north of Lake Naivasha to within ten 
miles of the east side of the Rift (there formed by the western foot of the Aberdares) . 
From this side of the Rift various collections possess birds of jacksoni type labelled 
Naivasha, II Polossat (at 7,000 ft. about fifty miles north of Naivasha), Rumuruti 
and Laikipia. These localities are all on the northern end of the massif of which the 
Aberdares form the crest and the local jacksoni appear to be separated by no ecological 
barrier from the Aberdares kikuyuensis. In particular, the forest at II Polossat is 
"mixed, with juniper, just the same as on the Aberdares" (J. G. Williams in litt.). 
Are then these birds allopatric? In any case the distances involved are very small 
and there is no sign of intergradation or hybridization between these two very 
different forms of Zosterops, both of which are essentially highland. 

The eastern birds of jacksoni type themselves form part of an interesting series. 
All the seven specimens from the area Naivasha-Rumuruti are a trifle yellower than 
typical jacksoni, but they are equally large. Sixty miles to the north-east, across a 
belt of dry thorn-bush, rather similar birds reappear on Gargues Mt. (about 8,800 ft.), 
the southern end of the Matthews Range, but they are smaller (four average 59-6 
against 62-0) and also duller, with a trifle less of both melanin and carotenoid. These 
birds may or may not extend along the whole seventy miles of the Matthews Range ; 
but on Mt. Nyiro (about 9,000 ft.), just detached from the northern end, the birds 
are slightly different. They are of the same ''difficult", generalized, type but rather 
larger than the Gargues birds and very variable in plumage. The seven Nyiro birds 



362 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

available average 62-5 mm. in wing compared with 59-6 in the Gargues. In plumage 
the Nyiro birds range from almost typical jacksoni to markedly duller and paler, 
like those of Gargues. (It is difficult to understand why Jackson and Sclater (1938) 
attributed Nyiro and Orr Valley specimens to the Abyssinian subspecies kaffensis, 
which has the carotenoid more golden and the forehead brighter and more defined.) 
To the east across eighty miles of thorn-bush, the jacksoni type of bird reappears, 
still further "attenuated", on the small volcano Marsabit, which rises to only 
5,500 ft. The forest is of a very dry type, with much Juniperus, and the rainfall 
exceptionally low for any sort of evergreen forest — only 32 inches at the meteoro- 
logical station 1,000 ft. below the summit. Twelve Marsabit birds are all dull- 
coloured and markedly small (wing 57-5) — but still quite different from the jubaensis 
(yet smaller and of a "dusty" pallor) of the surrounding thorn-bush. Thus we have 
a west-to-east cline of decreasing pigment and size on the three highlands, Laikipia- 
Gargues (Matthews Range)-Marsabit, all strongly isolated. 

As noted, the birds of Mt. Nyiro, off the north end of the Matthews Range, do not 
conform with the Laikipia-Marsabit trend: they are larger and not so dull-coloured. 
Their tendency in these respects is towards the Zosterops (kulalensis — see Note 17, 
Appendix 1) of the forest belt on Mt. Kulal (7,700 ft.), an old volcano forty miles to 
the north and also isolated by very dry country. The available specimens (from 
6,000-6,500 ft.) have the same wing-average, 62-5, as the Nyiro birds, but they are 
grey-bellied and perhaps also have rather longer tails and beaks. In other respects 
they resemble the Nyiro birds more closely than any others (and particularly than 
any of the Abyssinian birds, the nearest montane neighbours to the north). This 
situation finds an almost exact parallel in the relationship (described below) between 
the grey-bellied winifredae of the South Pare mountains and the yellow-bellied birds 
of the Usambaras a few miles to the south. And in fact morphologically the Kulal 
birds are to the South Pare birds just as the Nyiro are to the Usambara. The first 
two mentioned, which there is no reason to regard as closely related, show the same 
degree of convergence as the second pair. 

Reverting to the main eastern Kenya islands, for about 150 miles south from the 
southern end of kikuyuensis, near Nairobi, no montane Zosterops is known. Then, 
in the neighbourhood of Kilimanjaro and the Tanganyika border there are a number 
of forms. Birds resembling kikuyuensis in their broad eye-rings, but with less marked 
golden foreheads and less green on the underparts, appear in an arc of small mountains 
surrounding Kilimanjaro on the north, from the Mbulu Highlands and Mt. Hanang 
on the west, through Ketumbeine, Longido and the Chyulus to the North Pare Mts. 
(mbuluensis) — see Note 18, Appendix 1. A typical mbuluensis is known also from 
Essimingor Mt., of which more later. The very variable birds from the two extreme 
stations, Mbulu and North Pare, cannot, in series, be distinguished in plumage. 
(The fact that the latter average a little smaller, wing 60-4 compared with 62-5, is 
presumably at least in part because they come from altitudes averaging 1,000 ft. 
lower.) Yet between them intervenes Kilimanjaro, with a very distinctive Zosterops — 
eurycricota, with the largest size of eye-ring (up to 5 mm. broad), completely green 
forehead in nearly all specimens and, as a rule, much deep green and no bright yellow 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 363 

on the underparts. The difference between typical eurycricota and typical mbuluensis 
is striking, but individual variation is such that slight overlap occurs in each of the 
characters mentioned. 

Z. eurycricota extends westwards from Kilimanjaro, wherever there is forest, along 
the highland that connects Mt. Meru and Mondul Mt. with Essimingor, which is 
separated from Mbulu only by the twenty-five miles of the Rift Valley. Two speci- 
mens from Essimingor are exactly like Kilimanjaro birds, yet the third specimen 
known from this small mountain is the equally typical mbuluensis already mentioned. 
It is difficult to know how to interpret this. It could mean that eurycricota and 
mbuluensis belong to different species ; but perhaps the mbuluensis is a straggler 
from across the Rift Valley. A good sample of the Essimingor population is needed, 
especially as a basis of comparison for the future, in case one type of Zosterops is 
here replacing another. 

Birds like those of Kilimanjaro reappear in the attenuated forest on Lolkissale 
Mt. and the edge of the Lossogonoi plateau, fifty miles south of Kilimanjaro and cut 
off by arid thorn-bush. The only difference is that most of the Lossogonoi and 
Lolkissale specimens have a slight golden wash on the forehead, a feature that is 
very rare in the Kilimanjaro-Meru-Mondul population. 

About fifty miles east of Kilimanjaro, across, as it were, the Mbulu-Chyulu-North 
Pare arc, the grey-bellied birds of the Teita Hills (silvana) resemble the Kilimanjaro 
birds in their very large eye-rings and otherwise entirely green heads. Also they 
share with them the highest tail /wing ratio in East Africa, over 78. They differ in 
having a higher beak /wing ratio and more melanin generally — enough to make their 
throat greenish rather than yellowish — and a bright yellow edge to the "bend of the 
wing". This contrasts so sharply with the green of the rest of the upper parts that 
it is conspicuous in the field (M. E. W. North, in litt.) and at first strikes one as a 
peculiar feature. It is, however, only an exaggeration of the tendency, visible in a 
few eurycricota, for the tiny feathers on the bend of the wing to be slightly yellower 
than their neighbours. On the whole the Teita birds are undoubtedly more closely 
related to the Kilimanjaro than to any others. 

The Teita form is of special interest because it exists in exceptionally small 
numbers for a continental Zosterops. Fortunately for its survival, it is not entirely 
dependent on the few hundred acres of primary evergreen forest that remain in the 
patches. It has been observed in secondary Albizzia-Gummifera-Combretum 
woodland in the mountains, in flocks alongside, but not mingling with, flocks of the 
yellow savanna bird, flavilateralis (A. Forbes Watson, in litt.), and stomachs of both 
birds contained the same food. M. E. W. North estimates that the total population 
of silvana throughout the Teitas may be under 2,000 ; but similar birds inhabit the 
little cap of forest on Kasigau Mt., over fifty miles to the east and separated by arid 
thorn-bush. 

These grey-bellied Teita birds are only fifty miles from the Pare Mts. On the 
north end of this chain the yellow-bellied mbuluensis occurs, as already mentioned. 
Further south, the middle of the chain is lower and probably too dry for montane 
Zosterops, but towards the southern end, after an interval of some forty miles, 



364 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

montane forest reappears and is there inhabited by the last of the three grey-bellied 
birds of East Africa, winifredae. Apart from the belly-colour, these birds are rather 
intermediate between their northern neighbours, in North Pare, and their southern 
neighbours in the mountain forests of Usambara less than twenty miles away 
across the semi-desert Mkomasi gap. This might be due to clinal environmental 
influences of some sort, or, in view of the small distances, to gene-flow from both 
directions. The South Pare birds have carotenoid less golden than the North Pare, 
less prominent yellow on the forehead and tail /wing ratio lower — 73 compared with 
75. All these differences are tendencies towards the Usambara birds, which have 
slightly smaller eye-rings, less rich coloration and tail/wing ratio only 69. This 
Usambara type of bird extends down through Tanganyika Territory nearly to the 
Zambesi, and is further discussed in the section "Southern Tropical Africa". It is 
remarkable that on the East Usambara Mts., where evergreen forest extends down 
the seaward slope nearly to sea-level, the montane Zosterops in question stops at 
the edge of the plateau, at about 3,000 ft. Incidentally, no "savanna" Zosterops has 
been found round this seaward foot of the mountains or on the neighbouring coastal 
plain. 

Summary and classification 

Zosterops show sharper differentiation in this area than in any other. But among 
the montane forms extent of differentiation is not related to the extent of existing 
ecological barriers. 

The three grey-bellied forms each appear to be more closely allied to their nearest 
yellow-bellied neighbours than to any other Zosterops and not to be specifically 
distinct from them. 

On the same highland, at the north end of the Aberdares, jacksoni (from the west) 
and kikuyuensis (from the south) appear to meet allopatrically, without ecological 
barriers and without hybridization. This could mean that they belong to different 
species. The same applies to the occurrence of both mbuluensis and eurycricota on 
Essimingor but, partly because these forms show some overlap in characters, a 
different view is preferred. 

Apart from the belly-colour of the South Pare birds, the four montane populations 
of Usambara, South Pare, North Pare and Kilimanjaro, at first sight very dissimilar, 
form a (discontinuous) series of increasing pigmentation and size of eye-ring. This 
would suggest conspecificity (and the Usambara and Kilimanjaro birds, which look 
so different, have identical songs — see Appendix 2). Also, with mbuluensis as link, 
the Kilimanjaro eurycricota can be regarded as conspecific with the East Kenya 
kikuyuensis. 

Certain difficulties have now to be faced. The Usambara birds, just regarded as 
conspecific with the Kilimanjaro, are, as will be argued below, conspecific with the 
stuhlmanni of Lake Victoria. These are believed to intergrade, through the Yala 
population, with jacksoni. It follows that jacksoni would be conspecific with 
kikuyuensis, although, as noted above, the Aberdares situation suggests that they 
are not. A possible explanation is that here, as in the Great Tits, Parus major, of 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 365 

eastern Asia and other examples (Mayr, 1947), we have a meeting of branches of the 
same species so long separated as to be genetically incompatible. 

On the whole, I am prepared to treat all the montane Zoster ops as conspecific, 
retaining the names kulalensis, jacksoni (to cover the whole discontinuous cline from 
the West Kenya highlands through Laipikia and Gargues to Marsabit and Nyiro), 
kikuyuensis (East Kenya highlands), silvana (Teita), mbuluensis (including chyuluen- 
sis, for the arc Mbulu-North Pare), eurycricota (Kilimanjaro and outliers), winifredae 
(North Pare), stierlingi (Usambara ; see next section). The lowland birds, specifically 
distinct, are flavilateralis , intergrading with jubaensis (conspecific with abyssinica). 

CENTRAL AFRICA 

The area dealt with here is a highly mountainous strip about two hundred miles 
by six hundred, between 38 and 31 E., 3 N. and 6° S., on the eastern edge of the 
Congo Basin (see Map 7). Here the Zosterops show a wide range of variation in size 
and in general colour of plumage but not in eye-ring, forehead or belly colour. 
Although they include some of the deepest green birds in Africa, I regard them all 
as conspecific with the yellow West African senegalensis, which in the preceding 
section was traced south nearly to the Equator. 

Zosterops appear to be absent from the centre of the Congo basin 2 N.-5 S., 
cf. Chapin (1954), but are widespread in the east of it. Topographically this strip 
is dominated by the western (Albertine) Rift Valley and associated mountains, 
with the following main features, in succession from north to south : Lake Albert, 
Ruwenzori (17,000 ft.), Lake Edward, the Kivu volcanos (with forest belts up to 
over 10,000 ft.), Lake Kivu (at about 5,500ft.), Lake Tanganyika (at about 2,600 ft.). 
From north-west of Lake Edward to near the head of Lake Tanganyika the highland 
is continuous at 5,000 ft. or more, but Ruwenzori is isolated from this by the few 
miles of lower and drier country round Lake Edward. The highlands north-west of 
Lake Albert are more isolated (and the local Zosterops are so little represented in 
collections that they cannot be discussed). On their west side the Central African 
highlands decline rapidly into the fully lowland conditions of the Congo Basin with 
its equatorial forest climate. On the east, however, all the country southwards 
from Lake Albert, through Uganda and past the shores of Lake Victoria into 
Tanganyika Territory, lies at nearly 4,000 ft. or over. 

In almost the whole of this strip of Central Africa except the flats round Lake 
Albert the rainfall approaches 60 inches and considerable areas west of Lake Kivu 
and north-west of Lake Tanganyika, some below 3,000ft., receive over 80 inches. 
The wettest areas of all are the forested mountains, of which the chief are Ruwenzori 
and the Kivu volcanos. 1 The vegetation and appearance of the whole area have 
been graphically described, with photographs, by Chapin (1932). 

As previously stated, in Uganda the "yellow" Zosterops senegalensis appears to 
develop into the greener stuhlmanni as it approaches the shores of Lake Victoria. 

1 To the very local high humidity of these mountains the isohyets in Bultot (195°) probably do not 
do justice. 



366 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

Here it is a common bird wherever there are trees, at least from about Jinja round 
the northern and western shores of the lake and thence across western Uganda (but 
no Zosterops seems to have been collected on the other, drier, half of the lake shore) . 
The plumage of stuhlmanni is again of a very generalized type — diffuse yellow fore- 
head, medium eye-ring, greenish flanks contrasting with yellow centre of underparts. 
In fact the only feature that distinguishes these birds from those which are wide- 
spread on the mountains of Nyasaland and Tanganyika (and also the Imatong group 
on the Sudan border) is a certain warm tinge throughout the plumage. Uganda 
birds from between about 3,700 and 5,000 ft. have wings averaging 58-1, tail/wing 
ratio 71, dimensions which, like the plumage, are not distinctive. 

The Zosterops populations of islands in Lake Victoria would repay investigation. 
The only series is one of twelve kindly obtained for me by Mr. E. G. Rowe on 
Ukerewe Island, in the south-west of the lake (see Map 6). They have about the 
same-sized wing (58-8) as the mainland birds just described, but a higher tail-ratio 
(75), the Ukerewe eye-ring is larger, and the slightly warm tinge is absent from the 
plumage. In fact, lacking this tinge, in plumage the Ukerewe birds are separable 
from Imatong mountain birds — and indeed from birds on mountains round Lake 
Nyasa — only by slightly bigger eye-rings. Again, from Nkosi Is., the most exposed 
of the Sesse group in the north-west corner of the lake (see Map 7) , which is known 
to have faunal peculiarities (see Pitman, 1929), the single Zosterops available has a 
wing 4 mm. longer than that of any specimen from the opposite mainland and also 
an abnormally yellow head. 

Westwards these stuhlmanni extend to the wetter Ruwenzori massif, occupied by 
Zosterops up to at least 9,000 ft., and also to the low hot banks of Lake Albert. 
Between, there is a narrow corridor, mostly below about 3,000 ft., occupied by the 
Bwamba forest country (see van Someren and van Someren, 1949), leading north- 
west across the Semliki into the forested country of the upper Ituri, which declines 
to about 2,000 ft. In this highly diversified country on the western edge of stuhlmanni 
there is, as would be expected, much variation in Zosterops, which has been the 
subject of great differences in taxonomic opinion. There are adequate series only 
from Ruwenzori itself and from the upper Ituri (Stockholm Museum) and it is 
particularly unfortunate that no specimens seem to be known from localities linking 
the Ituri with the Ekibondo-Garamba area (already referred to under senegalensis 
as showing incipient divergence from that type) and with the smaller and more 
heavily pigmented birds atMedje, 150 miles to the north and the W.N.W. respectively. 

Eighteen birds are available from Bwamba and the low forest country to the 
west, from Kampi ya Wambuti to Beni, eighty miles to the south, which includes 
the type-locality of toroensis, Kitimba (which is not in present-day Toro). They 
are, as expected, all small, mean of wings 53*7, tails 36-9. 1 Also as expected, the 
birds are strongly pigmented, with plenty of both melanin and carotenoid, but they 
have little yellow on the forehead. On the whole they resemble the Medje birds more 

1 As Chapin (1954 : z ^7) ^ as noted, at Nganzi, near Beni, at 3,900 ft. at the western foot of Ruwenzori, 
he got a male with wing 61 alongside birds of toroensis size, The large bird might have been a straggler 
from the higher slopes, 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 367 

than any others and probably they intergrade with them over the intervening 
"uncollected" country. The Bwamba birds appear to be transitional between those 
of Uganda on the east (stuhlmanni) and the Ituri birds on the west (see Note 19, 
Appendix 1). 

There has been much difference of taxonomic opinion also about the Ruwenzori 
birds. I find that specimens from above 5,000 ft. (averaging 6,900 ft.) tend to have 
a little more melanin in the plumage and to be a little longer in wing and especially 
in tail than Uganda birds (altitude 4,000 ft.) to the east — wings 56-63 (mean 60-2) 
against 55-62-5 (mean 58-1), tails 41-49 (mean 44-5) against 37-44 (mean 41-3). 
(For the Ruwenzori mountain birds some authors have used the name scotti.) A 
series from the western foothills at Mutsora, 4,000 ft., averages the same size as 
Uganda birds at the same altitude on the east. 

In plumage both the higher-altitude Ruwenzori birds and those from the western 
foothills average a little darker than the Ituri birds (toroensis) and also than the 
Uganda birds. But the Ruwenzori population, with its generally strong melanin and 
"warm" carotenoid, cannot be separated in plumage from most of the slightly larger 
birds of Kivu (see below), a hundred miles to the south, on the other side of Lake 
Edward. In both the Kivu and Ruwenzori populations many of the birds have the 
melanin of the flanks extending towards the centre and clouding the whole underparts 
(as in virens of South Africa). This does not happen in Uganda birds, but at the 
same time there are individuals from the mountains that cannot be distinguished in 
plumage from Ituri or Uganda individuals. 

The identity between Kivu and Ruwenzori birds is noteworthy because the birds 
of the highlands just west of Lake Edward (see also Prigogine, 1953), which form an 
almost continuous connection above the 1,500 m. contour (approx. 5,000 ft.) between 
Ruwenzori and Kivu, differ slightly in being purer green, without the "warm" tinge. 
Moreover within the small area of these highlands, only some fifty miles by seventy, 
there is striking variation in size of Zoster ops with altitude. Birds from Lutunguru, 
on the western edge of the highland, and other localities between 1,400 and 1,500 m. 
(average 4,700 ft.) have mean of wing 57-1, tail 40-6, while birds from 1,600-2,700 m. 
(average 6,800 ft.) have wing 61-5, tail 46-6. Moreover the tail/wing ratio of the 
latter birds is 75 against only 71 for the former series — in accordance with the 
trend throughout the continent, cf . Part 3. A similar change in dimensions, correlated 
with altitude, takes place in the Kivu District. Birds from the volcanos average 
longer- winged and, more markedly, longer-tailed (62-5, 47*9) than those from nearer 
lake-level (60-9, 45*4). 

South-west of Lake Kivu and north-west of Lake Tanganyika lies an area highest 
in the east, dipping west into the Congo Basin at Kamituga, pioneered by Grauer 
but recently worked most usefully by Dr. A. Prigogine. The Zosterops (described 
as reichenowi Dubois) differ from those of the Lutunguru area, 150 miles to the north, 
only in having slightly more melanin (perhaps connected with the higher rainfall, 
80 against 60 inches). Like them, reichenowi lacks the warm tinge characteristic of 
Ruwenzori and most Kivu birds (as well as stuhlmanni) and indeed seems to have 
less carotenoid than any of the populations mentioned. It may be that further 



368 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

exploration will show a continuous narrow belt of these "colder^-coloured Zoster ops 
by-passing the Kivu population on the west (as adumbrated by the fact that Mulungu 
birds are a trifle greener) and intergrading with the small richly coloured birds of 
Bwamba-Ituri further north. 

The main interest of these southern birds, from Kamituga eastwards, which are 
all alike in plumage, is the change in their dimensions with altitude in a very short 
distance. The high-level birds, round about 7,000 ft., in the Muusi neighbourhood, 
are practically the same size as those at 6,800 ft. in the highlands west of Lake 
Edward — wings averaging 61-4, tails 45*3. But only forty miles to the west, round 
Kakanda-Kulundu, at 4,000 ft., the averages drop to 55-9 and 39-1 ; and twenty 
miles west again, round Kamituga and Utu, at about 3000 ft. (here the floor of the 
Congo Basin), the birds are smaller still, with wings only 53-1, tails 35-5. Further, 
a few geographically intermediate specimens fall perfectly on this size-cline. 

Summary 

While all the birds are regarded as conspecific with senegalensis, it seems worth 
while to retain the name toroensis for the small lowland birds N.W. of Ruwenzori 
and reichenowi for the " cold " green birds of the area N.W. of Lake Tanganyika. 
Ruwenzori and Kivu mountain birds are regarded as transitional between stuhl- 
manni of Uganda and reichenowi. Local variation in dimensions is great but 
consistent. It cannot be recognized nomenclatorially. 

SOUTHERN TROPICAL AFRICA 

This area (Map 8) extends from the southern limits of those dealt with in previous 
sections to the borders of South Africa. It contains a number of poorly differentiated 
Zosterops populations which differ in little but intensity of melanin, but have been 
allocated to one or other of two species on this criterion. 

I believe that this distinction cannot be maintained except for the lowland, dry- 
country form, flavilateralis, which extends south from Kenya, in a strip through the 
middle of Tanganyika to about as far as the line Tabor a-Morogoro. Between Tabora 
and Lake Victoria Zosterops seem to be absent ; and in southern Tanganyika 
Zosterops appear to be purely montane except for rare occurrences in the south-east 
(see Note 13, Appendix 1). 

The material of flavilateralis from Tanganyika does not suffice to define the local 
variation fully, but specimens from the moister country of Kondoa, Morogoro 
(Uluguru) and Usambara tend to be particularly green. They show in fact so much 
convergence towards the highland (forest-edge) birds of Uluguru and Usambara that 
some skins are not at first sight easy to allocate — even though the flavilateralis prove 
to have (B) melanization and the montane birds (A). The possibility of hybridiza- 
tion, for example at Lushoto (4,000 ft., Usambara) and Kibungo (700 ft. eastern foot 
of Ulugurus) whence particularly " difficult " skins come, cannot be excluded. 
However, I believe the best course is to regard these highland and lowland forms as 
belonging to different species, 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 



369 



Turning now to the Congo Basin, it appears, as already noted, that as far south 
as about 5 S. no Zosterops have been collected in the space of nearly 400 miles, 
between Luluabourg (in the Kasai country of the south-west) and the foot-hills of 
the mountains overlooking Lake Tanganyika in the east, or, to the south-west, 
Upemba (Map 8). But between the Kasai and the Transvaal, and from Angola to 



-IO*S 



5°5 



I5°S 



20°S 




Map 8. Southern Tropical Africa. 



the Indian Ocean, Zosterops with variable, but never highly distinctive, features are 
generally distributed. Some of them cannot be differentiated from the birds of the 
Imatong group of mountains far away on the Sudan-Uganda border, others differ 
from birds on the north and west of Lake Victoria only in slightly less "warmth" 
of tone, others again can be matched by abnormal West African individuals, others 
are extremely like the richly pigmented birds of Kivu and still others cannot be 
separated on plumage from individual South African birds. 

Taken as a whole, these southern tropical birds range in general colour of plumage 



37o VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

from predominantly yellow to predominantly green, and as there is a complete 
gradation the attempt to divide the populations between the species on this criterion 
alone, as is usually done, leads to difficulty. In dimensions all the populations, 
irrespective of colour, form one group of tail /wing ratios and beak /wing ratios (Text- 
figs. 4 and 8) and also when tail/wing ratios are plotted against altitude and against 
minimum temperatures (Text-figs. 6 and 7). (In the whole group of populations the 
tail/wing ratio is unusually constant in the face of changes in altitude and tempera- 
ture, and so is the beak-length with changing wing-length.) The main point for the 
taxonomist is that analyses of dimensions give no support to the view that more 
than one species of Zoster ops is present in the area under discussion. This view 
meets, however, with some difficulties on geographical and ecological grounds. 

Park populations are always, and dark individuals are nearly always, found at 
higher altitudes and where evergreen forest occurs, while the yellower birds usually 
occur in drier and lower country, dominated by deciduous woodland. But this 
ecological distinction is far from invariable. For example, R. H. Smithers has noted 
(in litt.) yellow birds in Southern Rhodesia in both "thorn veld" and evergreen 
growth, and Benson (1946a) has obtained such birds in the evergreen forest at 
Vumba (near Umtali, 5,000 ft.). Swynnerton, as his labels show, has collected yellow 
birds in the Chirinda and Chipete forests, but a dark bird in the Chimanimani ever- 
green bush half-way between these two localities. Again, some individuals on the 
border of Northern Rhodesia and the south-eastern Congo (Katanga) are so green 
that some workers have allocated them to virens (see Note 20, Appendix 1) — there 
are no mountains in the area, but some patches of evergreen growth too extensive 
to be called riparian (C. W. Benson in litt.). One is reminded of the dark individuals, 
formerly regarded as a separate form, phyllica, that crop up in the Bamenda high- 
lands (see p. 354). 

Lynes (1933) collected both yellower and greener birds (which he attributed to 
different species) in the same situations, the "edges of the forest-jungles" at 6,500 ft. 
in the Njombe District of Tanganyika Territory. Moreover, the Zoster ops of the 
Kungwe-Mahari mountains, on the east shore of Lake Tanganyika, are associated 
with evergreen forest (about 6,000 ft. upwards), yet when laid out with a long series 
of specimens from southern tropical Africa they fall into the yellower half. In 
Nyasaland the relict forest on some mountain tops (Mangoche, Dedza, Chongoni 
and Nchisi) is occupied by birds as yellow as those of the Rhodesian savannas, but 
greener birds occupy Mlanje, Cholo, Soche, Zomba, Ndirande and (greenest of all) 
Nyankhowa and the Nyika Plateau (Benson, 1945, 1953). No adequate meteorological 
data exist, but Benson (in litt.) is satisfied from his local experience that the moun- 
tains of the first group have a lower rainfall than the others. In none of the mountain 
forests where green birds have been collected has it been proved that a transition 
from yellower to greener birds takes place as the mountain is ascended and ever- 
green forest becomes the dominant vegetation ; and there seems no proof that yellow 
and green birds associate together. There is in fact no indication of such local clinal 
variation as previously noted on the slopes of the Imatong mountain group and 
elsewhere over more level country with a rapidly changing rainfall, as in Sierra 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 371 

Leone. The Nyasaland situation suggests at first the possibility that here we have 
ecological replacement of one species by another as in the instances given by Moreau 
(1948). For example, an isolated mountain forest, from which the typical montane 
woodpeckers (Mesopicos and Campethera) are absent, is inhabited by a species of 
Yungipicus that elsewhere is a savanna bird. Moreover, as will be shown in the 
next section ("South Africa"), a few hundred miles to the south-east of Nyasaland, 
in Zululand and southern Portuguese East Africa, yellow birds that are undoubtedly 
conspecific with, and have a distribution continuous with, those of Nyasaland appear 
to Jive as "good species" alongside, but ecologically separated from, green South 
African birds. 

These last closely resemble some of the darkest Nyasaland birds in plumage, but 
they differ from the Zululand yellow birds and from all the Nyasaland birds, both 
the yellow and the green, in having a higher tail /wing ratio, 75 against 67-71. Here 
we come up against the question of the value of this ratio as a taxonomic character 
(Part 3). On the other hand, as noted above, the yellow and the green birds of 
Nyasaland (and elsewhere in southern tropical Africa) do not differ in bodily propor- 
tions. It is a curious fact, however, that the dark birds tend to be unexpectedly 
small for the altitudes at which they are found. In particular, the dark Nyasaland 
birds (No. 64 in Appendix 3) have the same mean wing-length, 58-5 mm., as the 
local yellow birds (No. 63), with an average altitude 2,000 ft. lower and a climate 
certainly warmer. This is, of course, contrary to the trend otherwise general through- 
out Africa — and beyond — as discussed in Part 3. 

Thus arguments can be adduced for regarding the yellower and the greener birds 
in at least part of southern tropical Africa as belonging to two different species. 
Against this are the complete gradation that can be made up from birds from 
different localities and the fact that Benson (1953), who has an unrivalled knowledge 
of both forms in Nyasaland, can find no difference in their habits and their calls. On 
the whole it seems preferable to regard all the Zosterops of southern tropical Africa 
as belonging to the same species. 

The most north-westerly population in southern tropical Africa, namely, that in 
the Kasai district of the south-western Belgian Congo (kasaica), is the most dis- 
tinctive. They are the first Zosterops encountered after crossing the great forest 
from the north. The series from about 4 30' S., round Luebo and Luluabourg, 
consists of very small short-tailed birds (wing 52-5, tail 35*1), which conforms with 
their low altitude (ca. 2,000 ft. a.s.l.) and high minimum temperature (64 F.). On 
the whole they are more like the little Medje birds 700 miles away to the north-east, 
on the other side of the Congo Basin, than any others in Africa, but they are rather 
more olive throughout, with very narrow eye-rings and no clear yellow on the fore- 
head at all. They occupy country that, with an annual rainfall around 60 inches, is 
not so wet as that of Medje, but nevertheless the resemblance between these two 
populations, sundered by the forest, may be due to convergent evolution in similar 
equable climates, rather than to recent common ancestry. It is possible that a 
narrow belt of small birds remains to be discovered running eastwards from Lulua- 
bourg along the southern edge of the Congo Forest to intergrade clinally with the 



372 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

equally small, but greener, birds known from Kamituga (referred to in the section 
on Central Africa) at the foot of the eastern rim — themselves probably intergrading 
clinally to Medje. 

Towards the south-west, kasaica apparently extends at least to the north-eastern 
border of Angola — cf. the one imperfect specimen from Kasanga on the Kwango 
(Cuango) river. Thence, there is almost certainly intergradation with the birds of 
the highlands in the centre of Angola, which are much larger and much duller in 
plumage. The few specimens available from the intervening area Ndala Tando 
(= Vila Salazar)-Quicolungo-Canhoca (see Map 9), which is rather higher and drier 
country than that inhabited by kasaica, are already duller in plumage and much 
larger (wing 57-2, tail 39-0). 

Another 150 miles intervene to the south before another series of Zosterops is 
available from central Angola (Benguela Highlands) at 4,500 ft. upwards (Note 21, 
Appendix 3). This area has the highest rainfall in Angola 1 and it is therefore 
unexpected that the birds there (described as quanzae) are all duller in plumage 
than the Vila Salazar series. This may be connected with the fact that the Vila 
Salazar area is one of semi-evergreen forest (dominants Albizzia, Celtis and Ficus 
spp.), while the Benguela highlands, though having a higher rainfall, are occupied 
by savanna with much (deciduous) Brachystegia (Gossweiler and Mendonca, 1939), 
and with evergreen forest only in gullies (C. M. N. White in litt.). 

Also, the birds of the Benguela highlands are much larger (wing 61 -8) than those 
round Vila Salazar and in fact are the largest in southern tropical Africa (see Nos. 56 
et seq. in Appendix 3), although they come from an average altitude of only 5,500 ft. 
The explanation is presumably that, as data from ten local meteorological stations 
show, these birds experience abnormally low minima, averaging only 45 ° F. in the 
cool season — cf. the general size /temperature regression in Part 3. These large pale 
birds extend south-east at least to Vila Ponte (wing 64), but further down the 
Cubango (Rio Mbale) a specimen, though pale, is within the Rhodesian size-range 
(wing 58). 

Further collecting may bridge the present gap in Zosterops distribution in this 
part of Africa — nearly 400 miles to Dilolo in the east and 200 to Elephant Vley in 
the south — and show whether, as expected, there is a transition to the rather 
yellower and smaller birds (anderssoni) that occupy so much of the remainder of 
southern tropical Africa. There is at present no evidence that any such Zosterops 
occur further west in Angola, to the Atlantic, or further south in South West Africa, 
except along the northern border, or in Bechuanaland, but very similar birds are 
distributed through the Rhodesias, Nyasaland and Portuguese East Africa, right to 
the Indian Ocean, and also northwards in the south-eastern Belgian Congo to at 
least 8° S. The whole of this great area is, as already indicated, dominated by 

1 Cf. the isohyets in the Atlas de Portugal Ultramarino (Lisboa, 1948). They differ much in detail 
from those in Gossweiler and Mendonca (1939), but since they are presumably based on more data they 
are taken to be more nearly correct. As so often in climatological maps, the isohyets have had to be 
sketched in the Atlas from the records of widely scattered stations and it seems that in drawing them 
advantage has not been taken of the guidance orography can give. However, the main trends of the 
isohyets are no doubt somewhere near the truth. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 373 



IS"E. 



2&L 

BELGIAN 



CONGO 



IO°S. 



MOMBOLO 

(district) 



I5°S. 




Areas above 1,500 m. dotted. 
Map 9. Angola. 



zool. 4, 7. 



25 



374 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

deciduous woodland, though patches of evergreen forest occur, especially on the 
highest points. 

The yellowest populations of all are the most westerly, in western Northern 
Rhodesia. Equally yellow individuals occur as far east as Nyasaland (even at 6,000 ft. 
on the Nyika plateau), but most of the specimens from east of about 29 E. are a 
trifle greener than those from further west. The most easterly of all, in southern 
Portuguese East Africa (Mozambique), Zululand and the low-veld of the north- 
eastern Transvaal average a trifle greener still. Northwards through Portuguese 
East Africa it appears that the Zosterops generally become a trifle more richly 
pigmented, especially golden, a character that is best marked (but still only visible 
when series are compared) in the birds of the Songea highlands on the north-east 
side of Lake Nyasa. As the single bird collected between there and Indian Ocean, 
in the lowlands at Masasi, shows the same character, it may be that there is a con- 
tinuous, though extremely sparse, population over much of south-eastern Tanganyika 
Territory. These brightly coloured birds of south-eastern Tanganyika can, however, 
be matched by individuals from Nyasaland and Northern Rhodesia. Whether such 
birds extend far enough north to come in contact with the paleflavilateralis, hitherto 
not recorded south of the line Morogoro-Tabora, is unknown : and it would be 
interesting to know their relationships if they do so. 

In size the "yellow" populations of the Rhodesias, Nyasaland and also the south- 
eastern Congo, all around 4,000 ft., are remarkably constant, varying only from 
58-6 to 59-9 mm. in average wing and from 39-8 to 42-0 in tail (Nos. 59, 60, 62, 63 
in Appendix 3). A series from around 5,800 ft. in N. Rhodesia (No. 61) averages, 
as expected, larger (wing 61 mm., tail 42-5). By contrast, but again in accord with 
expectation, the birds from the coastal belt (below 1,000 ft.) along the Indian Ocean 
average only 56-6 mm. in wing and 39-2 in tail (No. 65). 

Returning to Northern Rhodesia, the variation in the Zosterops may now be 
traced north through the south-eastern Belgian Congo (Katanga). About 200 miles 
north of the territorial border, in the wooded savanna of the Upemba National 
Park (ca. 8° S. ; altitudes 4,500 ft .-6, 000 ft.) the birds are duller, apparently with a 
little more melanin and a little less carotenoid. In this they resemble the birds from 
the central Angola highlands (in fact Verheyen (1953) has used the same name, 
quanzae, for them) , but they are smaller — wing 59-9 against 61 -8. Similar populations 
occur on the west shore of Lake Tanganyika at Sambwe and Kabobo, the latter 
rather darker. Although there are great gaps east and west of Upemba, from which 
no specimens are known, it appears that we have a cline from the yellower birds 
(anderssoni) of the Rhodesias and Katanga, northwards through a stage deficient in 
both pigments to the deep green birds, with much melanin, in the highlands north- 
west of Lake Tanganyika (reichenowi — see section on " Central Africa "). It is 
curious that, as already mentioned, on the Kungwe mountains just across Lake 
Tanganyika the Zosterops are decidedly yellow. 

Turning now to the dark birds, the most richly pigmented of all are those in- 
habiting the Rungwe and Poroto Mts. at the north end of Lake Nyasa (see Note 22, 
Appendix 3), a small area that, with one station registering 100 inches a year, is 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 375 

about the wettest in southern tropical Africa. A few miles to the south, on the 
Nyika, Nyankhowa and Masuku Mts. of northern Nyasaland, the birds are nearly 
as richly coloured, with dark green flanks contrasting with the strong yellow centre 
of the underparts ; but on the other Nyasaland mountains, further to the south, 
none of the birds are so deeply pigmented and, as already noted, some are actually 
yellow. 

Facing the Poroto birds on the west, over 100 miles away across the Rukwa 
depression, the nearest montane Zosterops are those of the Ufipa highlands, and 
they, like those of the Kungwe-Mahari Mts., further north up the east side of Lake 
Tanganyika, are, as mentioned above, yellower. But eastwards and northwards 
from Rungwe the highlands are continuous to Njombe and Iringa, and all the 
Zosterops are dark, with a local variation that has greatly bothered taxonomists. It 
now appears that the birds on the average darkest, and most nearly resembling those 
of Rungwe, come from round Dabaga, on the top of the humid Uzungwe scarp that 
forms the seaward edge of the Iringa highlands. The birds that average next darkest 
are those of Njombe, while those from Iringa itself, which is not so high and humid 
as Dabaga and Njombe, are a little yellower. (For the nomenclatorial difficulties 
concerned see Note 23, Appendix 3.) 

Birds coloured like those of Iringa occupy the isolated mountains further north — 
Kiboriani, the Ngurus, Ulugurus and Usambaras — getting smaller as the altitude 
and latitude decrease and the minimum temperatures rise (populations 68-72, 
Appendix 3). As previously noted, the Zosterops of these Tanganyika mountains 
are partly islanded in the small, generally paler, Zosterops of the surrounding savanna 
which tend to converge in colour. These lowland birds to some extent penetrate the 
highlands — for example, reaching at least 4,000 ft. in the cultivated areas in the 
middle of the West Usambara plateau — but there is no evidence that the highland 
birds ever enter the lowland savanna. 

Summary 

The whole of the Zosterops inhabiting southern tropical Africa are regarded as 
belonging to the same species, senegalensis . It may be useful to retain the following 
subspecific names : kasaica for the small olive birds of the south-west Belgian 
Congo ; quanzae for the large dull birds of Central Angola ; anderssoni for the yellow 
birds from Northern Rhodesia to the Indian Ocean ; and stierlingi for the more 
richly pigmented birds of the Nyasaland and Tanganyika highlands. But the ranges 
of the last three cannot satisfactorily be delimited because intermediate populations 
are so prevalent and similar birds appear discontinuously, perhaps as a climatic 
response (convergence) . 

SOUTH AFRICA 

It would have been impossible to write even the present tentative account without 
much co-operation from ornithologists in South Africa, where I am especially in- 
debted to Mr. C. J. Skead for the Cape Province data and to Dr. R. M. Harwin for 
mapping the Transvaal situation. 



376 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES 




f 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 



377 





O 




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X 


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X 


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X 


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X 

X 


X 


X 







X 


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£ x 


® BRYAN S TON 

X x 




X 


GREEN BELLY 


O x 

o 

O 




° •<, o 




O 


PALE BELLY 








<3 


BOTH GPEEN ft PALE 








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BOTH VAAL & PALE 








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Map ii. Zosterops distribution in the southern Transvaal. 



378 VARIATION IN THE WESTERN ZOSTEROPIDaE (AVES) 

South of the Zambesi there are five main colour-forms of Zosterops, which have 
always been regarded as belonging to at least four different species. Their ranges 
and main characters are given in Maps 10 and n and Table 2 (see also Note 24, 
Appendix 1). For clarity the five forms are referred to in this discussion as "grey- 
bellies," "pale-bellies", "Vaals", "green-bellies" and"yellow-bellies , \ Briefly, grey- 
bellies differ from green-bellies only in lacking yellow pigment on the belly, but from 
pale bellies in deeper pigmentation of upper as well as under parts and in other 
characters. Vaals differ from pale-bellies only in having yellow throughout their 
underparts. Green-bellies differ from yellow-bellies (the anderssoni already discussed 
under "Southern Tropical Africa") in having more melanin throughout and individ- 
uals of them can be matched in colour with birds from the mountains of Nyasaland. 
All the purely South African forms have a high tail /wing ratio (75-79), contrasting 
with under 70 for the yellow -belly (anderssoni) that enters from the north. 

Table 2. — The main Characters and Distribution of South African Zosterops. 
N.B. — The subspecific names usually applied are inserted in brackets. 





Belly colour. 


Main area. 


Upper parts. 


(I) 


Grey (capensis 1 and atmorii) . 


Cape Province and Basutoland 


. Green 


(2) 


Whitish with chestnut wash on flanks 


Orange River Basin 


Paler and 




(pallida) 




greyer 


(3) 


Pale yellow with chestnut wash on 


Along lower Vaal R. (all locali- 


. As (2) 




flanks (vaalensis) 


ties shared with pallida) 




(4) 


Green, yellowish in middle (virens) . 


Eastern Cape, Natal, Zululand, 
Transvaal 


• As (1) 


(5) 


Yellow, green wash on flanks 


Zululand and N.E. Transvaal 


Yellower than 




(anderssoni) 


(thence northwards) 


(i)-(4) 



1 The most westerly grey-bellies (" capensis ") differ from all the others in this table in lacking any 
yellow over the lores (or over base of upper mandible). 

In South Africa the Zosterops arrangement differs from that elsewhere in Africa 
in that the colour-forms show considerable geographical overlap and even more 
ecological. All are limited to places where there are trees ; but, given that essential, 
they show a breadth of habitat tolerance that is unusual in African birds and is 
great enough to make it difficult to be sure what their original vegetational associa- 
tions were. 

(a) The grey-bellies occur from sea-level to over 7,000 ft. (Drakensberg). 
They inhabit all sorts of tree-growth, from the patches in the extremely dry 
Kamiesberg (south-east of the mouth of the Orange River) to the Knysna and 
Hogsback evergreen forests, including gardens and plantations of pine and 
eucalyptus (Skead). 

(b) Green-bellies are usually referred to as predominantly forest birds but, with 
as big an altitude-range as the grey-bellies and a wider geographical range, 
they inhabit if anything an even greater variety of arboreal vegetation — the 
coastal forests of the Eastern Cape, Natal and Zululand, the acacia-veld of 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 379 

the Natal uplands, the dry Transvaal high-veld, the gardens of Johannesburg 
and even the taller " bush " in the east of the Bechuanaland Protectorate. 

(c) Yellow-bellies are limited to the " low-veld ", the deciduous and pyro- 
phytic " bush " of the eastern Transvaal, Zululand, and Portuguese East Africa, 
thence spreading north and west across the Rhodesias. 

(d) Pale-bellies inhabit the southern half of South West Africa, the south- 
western Transvaal, the northern Cape and the whole of the Orange Free State — 
much of their area averaging less than 10 inches of rain a year. In the driest 
parts the birds seem to occur only in the valleys (especially along the Orange 
River), where there is some water, artificial or other (but there is no record of 
the birds drinking). 

(e) Vaal Zosterops are known only from localities on and near a short stretch 
of the lower Vaal River, from all of which pale-bellies are also known. 

As will be seen from Map 6, while Zosterops are generally distributed in South 
Africa, there are two areas where they seem at most very sporadic, (i) a wide belt 
in the western Cape Province from the Karroo northwards and (ii) most of South 
West Africa and the Bechuanaland Protectorate. The necessary trees exist scattered 
over much of the second area, but not in the Karroo. 

Local Variation 
Size generally 

As shown in Appendix 3 (populations 73-80), in each colour-type wings average 
longer in the cooler and /or higher areas, except that the green-bellies of the Natal- 
Cape lowlands average no larger than the Zululand birds that inhabit a slightly 
warmer climate (but by only about 2° F.). 

Grey-bellies 

East of about Port Elizabeth the grey-bellies average slightly yellower on the 
upper parts and throat and have some yellow on the forehead and lores. (This last 
feature corresponds with the amount of yellow on the heads of the green-belly 
population with which they interbreed — see below.) Enough specimens are not 
available, but the transition from the western form (capensis) to the slightly yellower 
eastern form (atmorii) appears to take place in a zone including Murraysburg, 
Graaff Reinet and Port Elizabeth, from all of which the specimens are intermediate in 
character. 

Well to the north-west of any other specimens of capensis two from the dry 
Kamiesberg are slightly duller above than other capensis and have flanks slightly 
redder. Both these features of their plumage adumbrate the characters of the pale- 
bellies, which are known from about seventy miles away on the north. However, 
something similar is shown by northern representatives of atmorii, further to the 
east, near Aliwal North (kindly communicated by Mr. P. A. Clancey). The plumage 
of these birds (and of one from Dewetsdorp) is duller throughout than that of the 
atmorii nearer the coast. On the flanks, however, these birds are a particularly pure 
grey, with no reddish, so that they can hardly owe their general divergence from 



3 8o VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

typical atmorii to an infusion of the local pale-belly characters. On the contrary, 
the dullness of the plumage appears to be correlated with the drier climate, for 
Dr. Auber has ascertained that a Kamiesberg capensis contains a lower proportion 
of (A) melanin and more (B) than a specimen from the more humid Knysna (see 
Part 2). 

Green-bellies 

These show much individual variation. Exceptionally dull specimens are scattered 
throughout the range, but every specimen from the driest part of the range (western 
Transvaal and eastern Bechuanaland — Note 25, Appendix 1) is slightly pale and 
dull, with the same divergence from the typical appearance as mentioned above for 
the dry-country representatives of capensis and atmorii. Moreover, the dull specimen 
(from Rustenburg) that has been tested shows a predominance of (B) melanin in 
contrast to the predominance of (A) melanin in a specimen from a humid Natal 
habitat (Dr. Auber). 

Pale-bellies 

Apart from size, no consistent local variation can be established, but the individual 
variation is very high. Besides occasional faint suffusion of yellow on the underparts, 
some specimens have the whole area between throat and vent nearly white, others 
almost entirely rufous. One from near Rustenburg, on the edge of the pale-belly 
range and some 250 miles from the nearest grey-bellies, has the underparts so grey 
that it would, if found south of the Orange River, probably have been cited as a 
hybrid atmorii X pallida. The occurrence of such a specimen, which must be of 
pure pale-belly stock, is important to bear in mind when considering another highly 
abnormal bird, shot at Dewetsdorp in the Orange Free State out of a flock composed 
partly of grey-bellies. This individual (Kafrrarian Mus. B2863) is predominantly 
warm russet below, becoming almost grey on the chest — a putative hybrid (see also 
Note 26, Appendix 1). 

General 

It will be observed that each of the colour-forms of extensive range in South Africa 
varies geographically in the same way. Each has its largest individuals in the 
coolest areas and its dry-country outliers less saturated in colour than the main 
populations in less arid climates. It is noteworthy that this consistent agreement 
with Bergmann's and Gloger's rules takes place within what are generally accepted 
as individual subspecies ; and the agreement with Gloger's holds good in the actual 
type of melanization so far as this has been tested. 

Overlap and Relationships of the Colour-forms 

Grey-bellies and green-bellies 

As shown in Map 10, there is a great overlap in the geographical distribution of 
these birds and, as already mentioned, no distinction in their habitat-preferences (or 
other behaviour). Some information (collated by Mr. Skead) is also available about 
their relative abundance in different localities. It can be only in general terms because 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 381 

colour of belly is difficult to see in such small restless birds and a formal census in 
an area where both occur is impossible. 

In the narrow belt about King William's Town, where grey- and green-bellies 
commonly interbreed (see below), they appear to be about equal in numbers. West- 
wards green-bellies diminish rapidly : at Queenstown, Somerset East and Cradock 
they are much in the minority, at Grahamstown and Port Elizabeth (less than 150 
miles from King William's Town) they are rare, and further west they are unknown. 
East of King William's Town the grey-bellies at first fall off rapidly, and green- 
bellies predominate heavily around Butterworth, Tsolo and Umtata, but further 
north, at the higher altitudes, in and on the borders of Basutoland, the two forms 
seem equally common again. Thus grey-bellies reassert themselves, as it were, in 
Basutoland and on the Natal slopes of the Drakensberg after apparently being some- 
what eclipsed by green-bellies nearer the coast on the south. In coastal Natal greys 
are rare, being known only from two specimens (Dargle and Durban — nearly 400 miles 
east of King William's Town). It is difficult to interpret these occurrences. The 
Durban specimen is too small to be a wanderer from the Basutoland population 
and, if a straggler, must have come along the coast from the south-west. 

Since Mr. Skead's field observations first suggested the relationships between the 
grey-bellies and the green-bellies he and his helpers have tried to record belly colour 
in every case of birds with nests or dependent young. The following results are all 
derived from the area King William's Town-Kei Road, where the two forms are 
constantly seen in the closest association : 

(1) Grey + green sharing incubation (of 3 eggs later destroyed) . 

(2) Grey + green with fledglings (belly-colour not ascertainable) . 
{3) Grey + green feeding one grey fledgling. 

(4) Grey + green with one grey fledgling. 

(5) Grey -j- green feeding three green nestlings. 

(6) Grey + green ,, three grey fledglings. 

(7) Green -j- green ,, three grey fledglings. 

(8) Green + green with three green fledglings. 

(9) Green -f- ? feeding three grey fledglings. 

(10) Green + ? ,, grey fledglings (? number). 

(11) Green + ? ,, one green fledgling. 

(In cases (9), (10) and (11) the second parent was not observed.) 

The above data are somewhat conflicting and the record that each bird, both 
parent and fledgling, is a typical green-belly or a typical grey-belly must be accepted 
subject to the fact that, as noted in Part 2, only a proportion of intermediates is 
obvious to the eye. (Indeed this happens even in field observations on the Carrion 
and Hoodie Crows — Mayr, 1947.) One "good" example of an intermediate Zoster ops 
has recently been collected by Skead ; its definitely grey flanks are separated by as 
marked yellow down the middle of the belly as ever occurs in the green-flanked 
type. Two other specimens from King William's Town and Newcastle have the 
flanks so dull greyish a green that they look somewhat intermediate, 



382 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

One definite conclusion from the field data given above is that the two green- 
bellied parents in (7) must have been heterozygous and that their fertility was normal, 
but no other genetical inferences can safely be drawn. Nor is the taxonomic problem 
settled. Bearing in mind also the geographical distribution, it seems that the grey- 
bellies and the green-bellies cannot be merely colour-phases. If the belt of inter- 
breeding of the two colour-types is really as narrow as that covered by the limited 
field-observations, then there might be a case for treating the two birds as species 
analogous to the Carrion and Hoodie Crows, but on the whole for the present they 
are best regarded as subspecifically related. 

Pale-bellies and grey-bellies 

Pale-bellies differ from the other South African Zosterops in having less saturated 
coloration — grey-green upper parts, paler yellow and wholly (B) melanin — presum- 
ably correlated with their drier environment. Their beaks tend to be more slender 
towards the tip, and in voice they differ slightly from both grey-bellies and green- 
bellies, which are alike. More important, their dimensions are peculiar in that their 
wings are abnormally short in relation to the low temperatures they experience 
(Part 3), though they share the high tail /wing ratio of other South African Zosterops. 

As will be seen from Map 10, in most of the area where pale-bellies and grey-bellies 
might be expected to overlap, i.e. in the south-western Cape Province, Zosterops of 
any sort have been very little recorded. The two forms have been collected together 
in only one locality, Dewetsdorp in the Orange Free State (out of the same flock ; 
Kaffrarian Museum), and the two specimens concerned each show a slight approach 
to the plumage characters of the other. The only other locality where the two forms 
have been found close together is Aliwal North (7 miles apart ; Mr. P. A. Clancey). 
As already noted, all the most northerly grey-bellies — those nearest to the pale-belly 
range — diverge slightly from the typical in the direction of pale-belly characters, 
but they can hardly be accepted as hybrids. Moreover, colour is not the only 
character separating the two types. 

Pale-bellies and Vaals 

Vaals are known only from the banks of the last 200 miles of the Vaal River and 
up to about thirty miles on either side (at Potchefstroom, Vredefort and perhaps 
Heidelberg), in localities where pale-bellies are recorded at the same season of the 
year, and mixed flocks are reported. The area has no notable ecological peculiarities 
(Dr. E. A. C. L. E. Schelpe in litt.). The relative abundance of the two forms would 
be impossible to assess reliably in the field. No biological information has been 
recorded about Vaals except by Plowes (1946 and in litt.) ; and it may be doubted 
whether any significant differences exist. 

Vaals have always been regarded as a distinct species (Z. vaalensis), but the five 
I have examined (only six specimens seem to exist in museums) show only one 
consistent difference from pale-bellies — they have a strong suffusion of yellow all 
over the underparts, which gives them a brighter yellow throat and a yellow belly 
with an isabelline, rather than a chestnut, wash on the flanks. Bearing in mind also 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 383 

the geography, specific separation seems unjustified and this is clinched by the 
existence of somewhat intermediate specimens. Two with slight yellow wash on a 
portion of the underparts come from Venterskroon and Bloemhof (Transvaal 
Museum) and one with yellow wash over the whole underparts from Rustenberg — 
100 miles from the Vaal. 

On the whole it seems best to regard "vaalensis" as a localized xanthochroic 
variety of pallida. Comparable cases are known. For example, the grass-warbler 
Cisticola juncidis, which has a vast range, from Spain to Australia and Natal, 
produces in a few square miles at the north end of Pemba Island, and apparently 
nowhere else, an erythristic form that interbreeds with typical birds (Pakenham, 
1937). 
Pale-bellies and green-bellies 

These two forms overlap in the southern Transvaal to the extent shown in Map 11. 
Critical information as to the extent to which they associate together in flocks, or 
interbreed, is lacking, but at Bryanston and at Hartebeestepoort both have been 
found in the breeding season. North of a line through Johannesburg greens occur, 
and apparently predominate, right into the dry country of the west, at least to 
Gaberones, over the Bechuanaland border, and the sources of the Great Marico, 
with ore locality of pale-bellies interpolated— Sterkstroom (Transvaal Museum). 
The furthest extension of green-bellies south-westwards is at Potchefstroom, where 
both pale-bellies and Vaals also occur. There is no indication of inter-breeding 
anywhere. 

Green-bellies and yellow-bellies 

At the northern end of their range the green birds from South Africa overlap the 
yellow birds of southern tropical Africa (anderssoni) in Zululand and the adjacent 
north-eastern Transvaal. This is the only case in South Africa where there is evidence 
of decisive ecological separation between two forms. In Zululand the green birds 
are said to be confined to patches of evergreen forest in the coastal acacia "bush" 
and to the forest on the Lebombo Hills (which do not exceed about 2,000 ft.), while 
the yellows occupy the surrounding more open vegetation (Mr. J. Vincent in litt.). 
It would be interesting to know whether the yellows and greens actually meet or 
associate anywhere, as, for example, the grey-bellies and green-bellies do constantly 
in the Cape Province. 

All the Zululand specimens available are either typical yellows or typical greens ; 
and it is important to remember that there is a difference in dimensions as well as 
in amount of melanin. The yellows average about 3 mm. shorter in wing than the 
greens and their tail /wing ratio is only 69 compared with 75 in the greens (and 75-79 
in all other South African Zoster ops). 

From Portuguese East Africa also both yellows and greens have been reported 
by Dr. A. A. Pinto (1953) and through his kindness I have been able to examine 
good series, supplemented by ecological data supplied by him. The only typical 
green ones are from three localities : 

(a) Namaacha, on the northern extension of the Lebombo Range, at about 



384 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

2,300 ft., where patches of evergreen forest occur — conditions exactly like those 
in which green birds occur in Zululand. 

(b) Umbelluzi, a locality practically at sea-level and devoid of evergreen forest, 
south-west of Lourengo Marquez. Yellow birds have also been collected here 
and greens would certainly not have been expected. The two greens (in the 
Smithsonian) were collected in July, and it has been suggested that they may 
be only winter- visitors to this untypical locality (Dr. Pinto, in litt. ; Mr. D. W. 
Lamm, in litt.). 

(c) Espungabera, ca. 2,800 ft. with evergreen forest patches, 400 miles to 
the north and only a few miles from Chirinda, over the border of Southern 
Rhodesia (see Map 8), has produced a typical yellow bird (wing 59, tail /wing 
ratio barely 70) and a typical green bird (wing 60, tail /wing ratio 75). As already 
noted, all the birds from the Rhodesian side of the border are all short-tailed 
and "yellow" except the one from Chimanimani that resembles the birds of the 
Nyasaland hill-forests. The Espungabera dark bird seems to be an isolated 
example of South African virens. 

Thus, on the evidence from Zululand, the yellow and the green birds would 
certainly be regarded as specifically different. The Portuguese East African distribu- 
tion is not so definitely in favour of this view, and moreover there are specimens 
whose plumage is intermediate. A male "yellow" from Umbelluzi (where, as noted, 
typical greens also occur) is darker and duller than the other Portuguese East 
African yellows ; two females from Boane and Lourenco Marquez are darker above 
than the other yellows, and one of them also has as much melanin below as a green- 
belly. Interbreeding has, however, not been observed. 

Taxonomic conclusions 
These are reached with hesitation and must be regarded as provisional : 

(a) Grey-bellies, capensis in the west intergrading with atmorii further east, are 
conspeciiic with green-bellies {virens). 

(b) This group is specifically distinct from yellow-bellies (anderssoni) . 

(c) Pale-bellies (pallida) form a monotypic species. (There is no reason at all to 
connect them with other forms elsewhere in Africa that lack yellow on the belly.) 

(d) Vaals ("vaalensis") are merely a colour-phase of pallida. 

PART 5 
INSULAR POPULATIONS 
THE GULF OF GUINEA ISLANDS 

The developments that have taken place in the Zosteropidae of the Gulf of Guinea 
are of special interest. Some of the birds concerned have so far differentiated that 
they have usually been regarded as worthy of separate specific and generic rank ; 
and, unlike what happens on the African continent generally, apparently complete 
sympatry of two forms of Zosteropidae occurs in several cases. The situation is 
sketched diagrammatically in Text-fig. n with mean wing and tail measurements, 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 



385 



There are four islands, all of volcanic origin and lying on an axis S.S.W.-N.N.E. 
that extends to Cameroon Mt. on the mainland. Annobon appears to be drier than 
the other (very wet) islands, but all still carry a good deal of primary forest. Their 
age is uncertain, but they are believed to be post-Miocene, the result of the tectonic 
activity that has affected the whole of Africa since the end of the long early-Tertiary 



63/13 




Fig. 11. The Zosteropidae of the Gulf of Guinea. 



386 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

calm. Exell (1944) has given a useful general account of the islands and Snow (1950) 
of the central pair, Sao Tome and Principe, with particular reference to bird- 
habitats. Amadon (1953) has recently published lists, with a comprehensive dis- 
cussion, of the Gulf of Guinea avifauna, in which, however, he has kindly left the 
Zosteropidae to me. 
The islands (Text-fig. 9) are, from north to south : 

Fernando Po, about 800 sq. miles, only 20 miles off-shore and rising to 9,300 ft. 

— 133 species of birds. 
Principe, 50 sq. miles, about 130 miles from Fernando Po and also from the 

mainland, and rising to 3,100 ft. — 30 species of birds. 
Sao Tome, 400 sq. miles, about 85 miles from Principe, 140 miles from the 

mainland, and rising to 6,600 ft. — 47 species of birds. 
Annobon, under 7 sq. miles, about no miles from Sao Tome, 210 miles from 

the mainland, and rising to 2,000 ft. — 9 species of birds. 

The three southern islands are typically "oceanic" in the biological sense and 
each is surrounded by water 6,000 ft. deep. Fernando Po is "continental", with sea 
only 200 ft. deep separating it from the mainland. There, little more than twenty 
miles from Fernando Po, stands Cameroon Mt., an isolated volcano rising to 13,300 ft., 
with which Fernando Po has the closest biological affinities. Indeed, for the purposes 
of the present discussion, Cameroon Mt. must be regarded as a Gulf of Guinea island. 
(The only other typically montane forest avifaunas in the western half of Africa 
are those on the mountains immediately to the north-east, Kupe, Manenguba and 
the Banso-Bamenda highlands, with which that of Cameroon Mt. has much in 
common.) 

The diversity of the avifauna on each island is no doubt partly a reflection of its 
size and variety of habitats, but also partly of its chances of receiving immigrants. 
Throughout the year the wind over this eastern part of the Gulf of Guinea is pre- 
dominantly S.S.W. (Meteorological Office, in litt.), thus favouring movement neither 
southwards down the chain Fernando Po-Principe-Sao Tome- Annobon, nor direct 
from the continent to any of the islands. On biological grounds, from a consideration 
of the insular avifaunas as a whole, Amadon (1953) has concluded that each of the 
three oceanic islands must to a large extent have received its birds independently 
from the mainland. 

Annobon is inhabited by only one member of the Zosteropidae, but Fernando Po, 
Principe, Sao Tome and Cameroon Mt. are each inhabited by two, one larger and 
with less yellow pigment than the other. Dimensions are given in Appendix 3 
(Part 2) and other salient details are summarized in Table 3 (see also Note 27, 
Appendix 3). Even though they show no striking modifications in beaks, there is, 
of course, a presumption that such sympatric pairs of closely related birds differ in 
their feeding habits, if not in their habitats (cf. Lack, 1944 ; Moreau, 1948). The 
larger birds of each of these Gulf of Guinea pairs differ widely between themselves, 
but they are all so different from normal Zosterops that they have been kept together 
in the genus Speirops (see Note 28, Appendix 1). It may be added that at least three 



VACATION IN THE WESTERN ZOSTEROPIDAE (AVES 



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388 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

of the Speirops and one of the insular Zosterops (Sao Tome) differ from all the con- 
tinental Zosterops examined by Dr. Auber in that melanization is not confined to 
the barbules but extends also to the barbs. 
What little is known of the ecology of the nine birds may be outlined as follows : 

(i) Cameroon Mt. at 6,000-9,000 ft. is inhabited by the brown endemic 
Speirops melanocephala, a bird which frequents the trees and shrubs in the 
montane grassland as well as clearings in the forest (Serle, 1950). Towards its 
lower limit this bird overlaps and associates with the smaller, yellow-green, 
Zosterops stenocricota, already discussed in Part 4. 

(2) On Fernando Po, the common Zosteropid is the Cameroon stenocricota (see 
Note 27, Appendix 1) ; the other species, the endemic brunnea, is so rare 
that only two specimens exist in museums and nothing is known of its ecology. 

(3) On Principe, the aberrant (grey) Zosteropid, Speirops leucophoea, is far 
bigger than its companion, the greenish ficedulina, but not bigger-beaked in the 
same proportion. No details are known of either bird, except that Snow (1950) 
found leucophoea in the lowlands, probably less abundant than it was first 
described to be in 1865, but yet more so than ficedulina, which he failed to see 
at all. (Correia collected some, however, in 1928 — Amadon, 1953.) 

(4) On Sao Tome, the situation is like that on Principe. The aberrant bird, 
Speirops lugubris, is the largest of all the western Zosteropidae. Its disparity 
in wing-length with the Zosterops of the island, feae, is greater than that between 
any other of the pairs under discussion ; but, as in Principe, the two Zosteropids 
show rather less disparity in beak- than in wing-length. Snow (1950) found 
lugubris only from about 3,000 ft., above which it was common in the forest ; 
but he tells me that, unlike what obtains on Principe, such vegetation is lacking 
at the lower levels, so that lugubris may not be typically montane. Snow did 
not see feae, though Correia was able to collect a series in 1928 (Amadon, 1953). 

(5) Annobon, so much smaller and less mountainous than the other islands, 
possesses only two passerines, a Paradise Flycatcher and the grey-green Z. 
griseovirescens (see Note 29, Appendix 1). It is the largest of the Gulf of Guinea 
Zosterops, smaller than the Speirops, but as big as most of the East African 
montane Zosterops. It was still plentiful throughout its wooded island when 
last reported in 1912 (by Boyd Alexander, in Bannerman, 1915). 

When the foregoing information is considered together with the dimensions in 
Part 2 of Appendix 3, and the other characters outlined in Table 3, several points 
of interest emerge, which are discussed below. 

Colour and pigments 

All the insular Zosteropids have tended to lose yellow pigment with the exception 
of the Fernando Po Zosterops. This shows at most only incipient divergence from 
the nearby Cameroon Zosterops, presumably because of its recent arrival on the 
island or to frequent interchange of populations (or to a combination of these two 
factors). 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 389 

As regards melanization, it is surprising that mainly (A)-type, i.e. black, pigment 
is present in the Speirops, which are predominantly brown or grey, and also in the 
Sao Tome Zoster ops, which is dull olive-grey-green. This colour is reproduced in the 
Principe Zosterops, which has not been examined microscopically, and also in the 
Annobon bird, but here its basis is (B) melanization (brown pigment) . Since Annobon 
is much the driest island, the difference in pigment type is correlated with climate in 
the expected way. On the other hand, it is entirely unexpected that the colour 
(as distinct from the pigment) of the Zosterops should be so similar on both the wet 
islands and the dry, and also so like the colour of the dry-lowland Zosterops of 
Abyssinia. It appears that the pale colour of the Sao Tome and Principe Zosterops 
cannot be a climatic adaptation. 

Colour of beak and legs also tends to be abnormal in the Gulf of Guinea birds. On 
the African continent all beaks are black except some of the Abyssinian, which are 
brown. All the Gulf of Guinea birds depart from this in various ways. And in having 
their legs and feet whitish or flesh-coloured instead of more or less grey the Cameroon 
and Sao Tome Speirops are distinguished from alLthe other Zosteropidae here 
reviewed. 

The white eye-ring that occurs complete, though in a variety of sizes, in every 
Zosterops on the African continent, is vestigial in the black-headed Speirops of 
Cameroon Mt. and the white-headed Speirops of Principe and is completely absent 
in the brown Speirops of Fernando Po. Apparently the eye-ring is lost fairly easily 
in Zosteropidae, as has occurred in uropygialis of one of the Kei Islands, alone among 
the Moris group inhabiting that area (Stresemann, 1931). In the Gulf of Guinea 
the white eye-ring is being lost not only in a white-headed bird, in which contrast 
is in any case minimized, but also in a black-headed bird, in which contrast would 
be particularly strong. 

Dimensions 

In size — as judged by wing-length — the insular forms that are aberrant in plumage 
have also all become abnormally large. The Principe and Sao Tome birds are both 
larger than any on the mainland of Africa. The only parallel is to be found in the 
most aberrant of the Pacific island birds, especially Rukia. Judging from the 
altitude /wing-length correlations on the continent (Text- fig. 1), the little-known 
Fernando Po bird is also abnormally large unless it is confined to the highest 
part of the 9,000-foot peak. Each of these aberrant birds presumably belongs to 
stock that has been longer on the island concerned than the less aberrant local 
Zosterops ; and Amadon (1953) has already noted that the tendency of Gulf of 
Guinea insular birds to be big is especially marked in presumptive first-colonists 
such as these Speirops are. 

From south to north the four Speirops form a series of diminishing size — Sao 
Tome 74 mm., Principe 68, Fernando Po 65, Cameroon Mt. 63. No general reason 
can be suggested for this. As shown in Part 3, on the continent wing-length is closely 
correlated with both altitude and temperature. Lack of meteorological records and 
ignorance of the average altitude of the available specimens of each Gulf of Guinea 

ZOOL. 4, 7. 26 



39© VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 

form make it impossible to test the temperature correlation for these birds. It is, 
however, most unlikely that it would hold good, because the Cameroon Speirops, 
which is the smallest of the birds in question, comes from much the highest average 
altitude (8,000 ft.). The other stations are both more maritime and nearer the 
equator ; and there is no reason to suppose that the insular Zosteropids in question 
experience lower temperatures than the high-altitude birds of Cameroon Mt. I 
conclude that the large sizes attained by the Speirops are due to local biological 
causes and that there is no general reason for their arrangement in a series of 
decreasing wing-length northwards. 

In contrast to the Speirops, except on Annobon none of the Zosterops diverges 
obviously in size from expectation (though again the data do not suffice for correlating 
wing-length with temperature). On Annobon, which does not rise above 2,000 ft., 
and which contains only one passerine besides the Zosterops, the latter is larger than 
expected. But on each "island" possessing two Zosteropids the less aberrant bird is 
the smaller, to a very variable degree. The difference in mean wing-length between 
Speirops and Zosterops is 19 mm. on Sao Tome, 16 on Principe and only 9 mm. on 
Cameroon Mt. and Fernando Po. 

Tail/wing ratios vary remarkably. At one extreme the Fernando Po Speirops 
appears to have a longer tail in proportion to wing (ratio over 82) than any other 
population of the western Zosteropidae (though individual South African birds 
exceed 80). At the other extreme the Sao Tome Speirops has an abnormally short 
tail (ratio 61 -8), equalled in this respect only by the Zosterops of the opposite main- 
land (as exemplified by those of Cameroon, with ratio 61 -6). The insular Zosterops 
show a similar lack of uniformity. The Fernando Po bird is, like the Cameroon, 
abnormally short-tailed (62-8) by continental standards. The Principe and Sao 
Tome birds have much higher, more "normal", ratios, 71-7 and 67*2 respectively, 
while the Annobon bird with ratio 77-6 is at the top end of the continental range. 
Thus there is no reason to regard the short tail of the Sao Tome Speirops as evidence 
of particularly close affinity with the short-tailed but otherwise normal Zosterops of 
the neighbouring mainland. In fact, as discussed in Part 3, tail/wing ratio is of 
uncertain taxonomic significance. 

The beak /wing ratio varies in the nine forms under discussion between 21% and 
26%. The insular Zosterops have longer beaks in proportion to their wings than the 
insular Speirops (this does not apply to the Cameroon Mt. populations). In fact, 
against a mean beak/wing ratio of 22-7 for the continent of Africa as a whole, the 
mean beak/wing ratio of the Speirops in the Gulf of Guinea islands is only 21-3, while 
that of Zosterops is 24-5. Thus the Zosterops, but not the Speirops, accord with Ama- 
don's (1953) generalization (based on other species) that insular birds tend to have 
larger beaks than their nearest relatives on the mainland. This might suggest that 
the Speirops, in the course of their evolution to their present stage of abnormality, 
passed through a phase in which their beak /wing ratio rose above that of the 
continental birds but that later, with increasing body-size, the process was reversed. 
On this hypothesis the Annobon bird is in the intermediate stage, with beak /wing 
ratio still high but body-size becoming abnormally large. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 391 

General 

The Zosteropids of the Gulf of Guinea must have been derived originally from 
the opposite mainland, in at least some of the islands by two distinct colonizations. 
But which of the existing forms were evolved in situ from colonists direct from the 
mainland and which were derived from inter-island invasion cannot be determined. 
Any discussion could be little more than speculative, especially as the characters 
of the ancestral mainland Zosterops can only be surmised. 

A main general change in all the insular birds is the loss of yellow pigment ; and 
it is difficult to suggest why this should be so, especially since, as has been shown, 
plumage colour is not correlated with climate. 

The several Speirops differ from each other and from "normal" Zosterops in so 
many characters and to such varying degrees that it is impossible to decide which is 
the most primitive or the most "specialized". Those of Cameroon Mt. and Sao 
Tome are sufficiently alike to be regarded as conspecific, and this although two islands, 
each with a very different Speirops, intervene geographically. The Sao Tome bird 
has not lost eye-ring and yellow pigment to the same extent as the Cameroon bird, 
which suggests that the former is the nearer to the ancestral stock, and hence that 
the Cameroon bird is an invader from an island. On the other hand, the whitish on 
the forehead and throat of the Cameroon bird, which is lacking in the Sao Tome, 
is adumbrated in those numerous "normal" Zosterops which have yellow on the 
forehead and throat contrasting with a generally green head. Hence in head-pattern 
the Cameroon bird may be regarded as retaining primitive characters that the Sao 
Tome bird has lost. In any case it seems best to recognize the closeness of resemblance 
between the Cameroon and the Sao Tome Speirops by treating them as conspecific. 
The Principe Speirops, with its very pallid foreparts, and the Fernando Po bird 
have evolved on different lines from the other two. 

The Zosterops of Principe and Sao Tome are so alike that one is more likely to 
have been derived from the other rather than both evolved, convergently, from 
independent colonists from the mainland. In general colour the Annobon Zosterops 
is so like these two that it has sometimes been regarded as conspecific ; but it is 
sufficiently different in size and proportions for this to seem undesirable. 

Hence we get the following classification : Zosterops senegalensis stenocricota on 
Fernando Po, Z. ficedulina ficedulina on Principe, Z. ficedulina feae on Sao Tome, 
Z. griseovirescens on Annobon. Speirops lugubris melanocephala on Cameroon Mt., 
5. lugubris lugubris on Sao Tome, S. brunnea on Fernando Po and 5. leucophoea on 
Principe. 

THE ISLANDS OF THE INDIAN OCEAN 

Zosterops are widely distributed in the Indian Ocean and belong to at least six 
species. Most of the islands fall into two groups (see Map 12), those inhabited by 
Zosterops being denoted in the following paragraphs by an asterisk. 

(a) Low limestone islands, mostly atolls, of which there are a great many, especially 
in the east, where the Laccadive*, Maldive and Chagos archipelagos are separated 



392 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES 




CHA COS 



.» CAR G ADOS 



A © RODRIGUEZ 

MAURITIUS 
REUNION 



Map 12. The Indian Ocean. 



VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 



393 



by 1,200 miles of uninterrupted sea from the Mascarenes. Other coral islands form 
the Cargados, the Amirantes, other outliers of the Seychelles and, nearer Madagascar, 
the Aldabra group*, Gloriosa*, and, in the Mozambique Channel, Juan de Nova 
(17 4' S., 47 43' E.) and Europa Island* 22° 20' S., 40 20' E.). None of this class 
of island has an area of more than a very few square miles and most of them are 
only a few hundred acres. The natural vegetation appears to have been scrub jungle 
and much of it has been replaced by coconuts. 



Table 4. — Main Colour Characters of Indian Ocean Zosterops. 

N.B. — All have a small eye-ring. 



Islands. 


Upper parts. 


Yellow on head. 


Underparts. 


Madagascar group : 








(1) Madagascar 1 maderaspatana 


Yellow-green 1 


None. 


Whitish more or 

less washed 

brownish grey 


(2) Gloriosa .... 


As the yellowest 
of (1) 


>» 


As (1) 


(3) Cosmoledo 


? Paler and less 
yellow than (1) 


»» 


• >» 


(4) Europa .... 


As (1) 


»» 


• >» 


(5) Aldabra aldabrensis . 


Yellower than 


Variable on lores 






(2) 


and forehead 




(6) Laccadives 


Yellower than 


As ( 5 ) 


As (1) but 




any above 




clearer 


Comoros : 








(8) Grand Comoro comorensis . 


As (2) 


On lores and 
forehead 


As (1) 


(7) Anjouan anjouanensis 


Slightly yellower . 
than (1) 


On lores 


• >» 


(9) Grand Comoro kirki . 


Yellow green 


On lores and 


Golden-yellow, 






forehead 


green on flanks 


( 1 0) Grand Comoro mouroniensis 


Greener than (9) 


On lores 


Greenish yellow 


(n) Mayotte mayottensis . 


Greenish-yellow . 


On lores and 


Golden yellow, 






forehead 


reddish at sides 


Seychelles : 








(12) Marianne semiflava . 


Slightly yellower . 


Less than (11) 


Brighter than 




than (11) 




(11) 


(13) Mahe modesta . 


Grey-brown 
tinged olive 


None 


. Pale grey-brown 



Mascarenes : 
See Table 5 

1 Marked local variation, the birds being darkest in the wettest areas, yellowest in the driest. 



394 VARIATION IN THE WESTERN ZOSTEROPIDAE (AVES) 




X Grey belly O Whole plumage greyish 
+ Yellow belly